Full text of "Malacologia"

See other formats

5 4 4 ‘ as у à в i H + : y 23 > À 4 ñ 
2 Si $ : Ка р is р 
7 ‘ aie DAME | A } . у ыы Ne 
ER eh 11 + р 4 q ИЗ < . р | x val : : 164 ale 
1, 7 rivers ; a 4: ae oe 
5 2 4 9 > cata Ait : A E > 
: т 4 $ В 4 A 7 El 4 0 RD EU pat Н + Den iy 
: eT A lan 4 a IE “ail 5 £ Hy AS à 
4 a 174 я bs 2 Чан = 5 ie Ма Fu = Speke £ 2e 1 И ELA 
i tty) gt ey SERA AN ANS : : Au A om he vas 
7 : : N an . Spray tn! } Be dam À 4 Sup PAR 


Kurs 
Viva 
jus 


Ways 
А 
Че, 
р 
ye 


4 
“ 
р 


pego 
pls 


LA 
ue: 
due 


a an 


‘i ver DENE i 1 ¥ : ; 4 Kann a dadas 


mapa 
REN 
rept yee 


TA DEA 
SN DENE: 


a 


DEN EN 


и N 
RA 


И 
Фот: 
АЕ 
АА Uy: 

A 

Soe} 


AVES LE ENTRE 
НЕ 
ить 

14 
Е 4 
Азы: 


RE REN 


» } : à 44 ? ind) 
E у if the añ 


а 


He ya 

ted ‘ 

гаи 

WARIS atado ón 
РИН 

и 


Fer ee Ана, 
Dre pty sy ata 


apt yee 


HR era. 

‘ ль 
еее 
Eur, 


CHEN 
VE 
aan 


Magan ja 


4 


ces 


a 


sty pal IIA 


FA Ben pacs 


shed ay aoe 
tte 
UNA ge 


dann 
an ие, 


RTC 


Пауза 


же: 
2 


nt 


LATE 


4124 
usa 


QUE 


MARTINA OO 


7 


аи ву. 


14 


Lips 
Beyer 


usa 
any 
[Au rays 
inte REP 
sas 
CRUE 


Mágina 


rate 


A 
O) 


asain i 


vere ey 
Ry tye, 
a 


Anus, 
uch 
PTIT 

anges 


era 

ae EN ON 
PA RO 
ALI 


MULA UL e 
ti 
RTE TEEN 


ИИ 


PIE IR EU eye 
tle peu tye 
AO DAB ody 

oot 
4 


Чье 
tas 


мора: EP take 


CRA 


sO) tn be 
ХИ DS EP 
DEN 


14,8 


ba 
vey 


MMA EE 


Tekan 


says 


ES 


Leet 


Зе 
TORE Ea ЗНА A 


tin 


а вре 


We ea pee 


era RAR YS 
AVATARS 
KATZE 


AA 
OA 
ES 


RA 
th votes 
pra 
PRE 
sea) ter 
аа 

ре 


Let IQ enr 


ааа 
une 


«на 
Аля. 

т 

«А 


Feu 

eee gage 
ney Oye pat 
Aue eee 


ES 


Pones, 
Pa do 
A 


NE COR АЯ 


А ве 


Ag eas у2 


esta ate venus 


А A 


4 DET 
PUR PUR 


EME BEEN a yet es 


249 
RYE eRe Gay teeta 
en 


4, 
24d 


DAA 


pad Do TN 
apra 
Ma На, вре 
yes end. 1$ yoy 
DATE y 
pS Ya ey 


bear 


y BY IS A O) 
EUR Hi het ped, 
ay 
р CET DETENTE ET ER RIRE 
à FU АА 
PAUSED dehy удаль лю DEE 
Icy Se MEAS ra Gp 
PEE DET зи ре 
E 
AI 


A 
ua Ap 


OR 
pa 


MEDINA Cott £7 
age 
Vas etary ii) 
Aaa 


AT he's 


а 


gg 
y > 


MALACOLOGIA 
http://malacologia.fmnh.org 


Worlds Leading Malacological Journal 
Journal Impact Factor 1.48 (2006) 


EDITOR-IN-CHIEF 
GEORGE M. DAVIS 


Editorial Office Business & Subscription Office 
Malacologia Malacologia 
| РО. Box 1222 P.O. Box 385 
West Falmouth, MA 02574-1222 Haddonfield, NJ 08033-0309 
georgedavis99@hotmail.com malacolog@jersey.net 
Copy Editor Associate Editor 
EUGENE COAN JOHN B.BURCH 
California Academy of Sciences University of Michigan 
San Francisco, CA Ann Arbor, MI 
gene.coan@sierraclub.org jbburch@umich.edu 
Managing Editor Composition Editor 
CARYL HESTERMAN CLAUDIA WILKE 
Haddonfield, NJ Giessen, Germany 
malacolog@jersey.net claudiawilke@hotmail.com 
Graphics Editor 


THOMAS WILKE 
Justus Liebig University 
Giessen, Germany 
tom.wilke@allzool.bio.uni-giessen.de 


MALACOLOGIA is published by the INSTITUTE OF MALACOLOGY 
Board of Directors: 


President JOHN BURCH 
ALAN KOHN University of Michigan, Ann Arbor 


University of Washington, Seattle DIARMAID © FOIGHIL 
University of Michigan, Ann Arbor 


CAROLE S. HICKMAN 
University of California, Berkeley 


PAULA M. MIKKELSEN 
Paleontological Research Institution, Ithaca 


Vice President 
RUDIGER BIELER 
Field Museum, Chicago 


Secretary and Treasurer 
GEORGE M. DAVIS SHI-KUEI WU 
West Falmouth University of Colorado Museum, Boulder 


Emeritus Founding Consultant 
J FRANCES ALLEN 


Copyright © 2008 by the Institute of Malacology 
ISSN: 0076-2997 | 


J. A. ALLEN 
Marine Biological Station 
Millport, United Kingdom 


T. ASAMI 
Shinshu University 
Matsumo, Japan 


T. BACKELJAU 
University of Antwerp 
Antwerp, Belgium 


B. BAUR 
Universitat Basel 
Basel, Switzerland 


Е. Е BINDER 
Muséum d'Histoire Naturelle 
Geneve, Switzerland 


D. BLAIR 
James Cook University 
Townsville, Australia 


P. BOUCHET 


Muséum National d’Histoire Naturelle 


Paris, France 


J. BUCKLAND-NICKS 
St Francis Xavier University 


Antigonish, Nova Scotia, Canada 


P. CALOW 
University of Sheffield 
Sheffield, United Kingdom 


К. А. D. CAMERON 
University of Sheffield 
Sheffield, United Kingdom 


J. G CARTER 
University of North Carolina 
Chapel Hill, NC 


B. C. CLARKE 
University of Nottingham 
Nottingham, United Kingdom 


T. COLLINS 
Florida International University 
Miami, FL 


К. Н. COWIE 
University of Hawaii 
Honolulu, HI 


2008 
EDITORIAL BOARD 


SY HSONIAY 


JUN 20 cuvo 
LIBRARIES 


G. CUEZZO 
Universidad Nacional de Tucumán 
Tucumán, Argentina 


B. DAYRAT 


University of California 
Merced, CA 


SMIT HSON/A 
JUN 20 zu0g ` 
LIBRARIES 


Ко ВО, JE 
College of Charleston 
Charleston, SC 


C. J. DUNCAN 
University of Liverpool 
Liverpool, United Kingdom 


G. DUSSART 
Canterbury Christ Church University College 
Kent, United Kingdom 


D. J. EERNISSE 
California State University Fullerton 
Fullerton, CA 


D. GEIGER 
Santa Barbara Museum of Natural History 
Santa Barbara, CA 


G. GIRIBET 
Harvard University 
Cambridge, MA 


E. GITTENBERGER 
Rijksmuseum van Natuurlijke Historie 
Leiden, Netherlands 


Е. GIUSTI 
Universita di Siena 
Siena, Italy 


M. GLAUBRECHT 
Museum of Natural History Berlin 
Berlin, Germany 


A. N. GOLIKOV 
Zoological Institute 
St. Petersburg, Russia 


D. GRAFF 


The Academy of Natural Sciences 
Philadelphia, PA 


T. HABE 


Tokai University 
Shimizu, Japan 


R. Т. HANLON J. LEONARD 


Marine Biological Laboratory University of California 
Woods Hole, MA Santa Cruz, CA 

E. HARPER C. LYDEARD 

University of Cambridge University of Alabama 
Cambridge, United Kingdom Tuscaloosa, AL 

G. HASZPRUNAR H. K. MIENIS 
Zoologische Staatssammlung München Hebrew University of Jerusalem 
München, Germany Jerusalem, Israel 

J. M. HEALY J. J. MURRAY, Jr. 
Queensland Museum University of Virginia 
South Brisbane, Australia Charlottesville, VA 

D. M. HILLIS 

University of Texas R. NATARAJAN 

Austin, TX Marine Biological Station 


Porto Novo, India 


К. Е. HOAGLAND 
|. М. Systems Group 


Rockville, МА J. OKLAND 
University of Oslo 
W. HOEH Oslo, Norway 
Kent State University 
Kent, OH T. OKUTANI 
University of Fisherie 
B. HUBENDICK Tokyo fae Lis 
Naturhistoriska Museet i 
Goteborg, Sweden VW. L. PARAENSE 
$ HUNT Instituto Oswaldo Cruz 


Rio de Janeiro, Brazil 
University of Central Lancashire | [ores 


Lancashire, United Kingdom 


J: J. PARODIZ 
Carnegie Museum of Natural History 


C. ITUARTE 
Museo Argentino de Ciencias Naturales 


Buenos Aires, Argentina Pittsburgh, PA 


R. JANSSEN 
Forschungsinstitut Senckenberg 
Frankfurt am Main, Germany 


G. PAULEY 
Florida Museum of Natural History 
Gainesville, FL 


M. S. JOHNSON 
University of Western Australia 
Crawley, Australia 


R. PIPE 
The Marine Biological Association 
Plymouth, United Kingdom 


G. KAMANEV 

Institute of Marine Biology, Russian Academy of a ie alas 

Science Ecole Pratique des Hautes Etudes 
Vladivostok, Russia Perpignan Cedex, France 

У. KANTOR М.Е. PONDER 

Russian Academy of Sciences Australian Museum 

Moscow, Russia Sydney, Australia 

R. N. KILBURN T. RAWLINGS 

Natal Museum Cape Breton University 


Pietermaritzburg, South Africa Nova Scotia , Canada 


О. ©. REID 
The Natural History Museum 
London, United Kingdom 


T. SASAKI 
University Museum, University of Tokyo 
Tokyo, Japan 


$. С. SEGERSTRÄLE 
Institute of Marine Research 
Helsinki, Finland 


L. К. L. SIMONE 
Museu de Zoologia da Universidade 
de Sao Paulo, Brazil 


A. STANCYKOWSKA 
Siedice, Poland 


F. STARMUHLNER 


Zoologisches Institut der Universitat Wien 
Wien, Austria 


J. STUARDO 
Universidad de Concepcion 
Valparaiso, Chile 


P. TATTERSFIELD 
Tideswell, Buxton 
United Kingdom 


C. THIRIOT 
University Pierre et Marie Curie 
Paris, France 


5. TILLER 
Muséum National d'Histoire Naturelle 
Paris, France 


A. VALDES 
California State Polytechnic University 
Pomona, CA 


J. À. M. VAN DEN BIGGELAAR 
State University of Utrecht 
Utrecht, Netherlands 


N. H. VERDONK 
Rijksuniversiteit 
Utrecht, Netherlands 


J. VOIGHT 
Field Museum Natural History 
Chicago, IL 


J. VOLTZOW 
University of Scranton 
Scranton, PA 


H. WAGELE 
Ruhr-Universitat Bochum 
Bochum, Germany 


A. WAREN 
Museum of Natural History 
Stockholm, Sweden 


M. WU 
Hebei University 
Hebei, China 


Арн А meen Pa Г | 7 | 


ive a, Ma De oe 


t 


a "di 
pi 


bis A 
qe u m 


he: я 


% 


> Y a enr | NESTS RATE A 
: р = ie Tien er “ : 
fica MUA E E e hi LL ы ho ot ay N er м 
aa р 
Mr “beaks va 


i 
CRE re 
i wi Dit И 


RES 


gs 
a 
ul 4: 
ы м EE И > 
Un ER вы $ 
E = LL 


A В р u a qn Es 
DEC. 


Е Hut + г 
К Sent no 209 
LR ke thin atid 


=; A } 
GUIBIMARATE A | 
| 


ou arre & 
2. Loan AA 
u ei ES E 


ae "sir 


ee Re Mer; RER. 
ee ee ld feet 2 


Mh >>. ны! nf nah x Se HER iR Cats: 22 


2 E 


ae BR: el, Be: ‘ 
O dr e OE 


a 1 Korn mes, = cz en Y aan т | 


5 | уф Eu Baar es RARE El D 
ER age de ae din RR PO 
1 

i 


+ tr 5 wee 


E 


AS A | et oi 5 " меж, u 
Me rer cd oh a а ahi ee DER E ras DRE 
= of и à a = 7 Ya x sr + sl = Pe a a 


ы - Pr 
rea та * » $ 3 ep "a (m 
« a 5 


y ay sh are Sc hed sa 7 ai” 
> вы de e y 2 Br o z г и cua a 5 


2 x Pee eee Pal ke Oe: A 
+ : he moi E re Pee a A = ск. 
q a ae el 4 

RS A Ц pa | у р | 


Cee Е = < f ge ok = “+ va. 
= у 4 ES x 

rsa? 2 : 

A 


LR: ы a = на PRET. 


Sue Po PT CR et pa SE = = + a Ku See de LOGE. q 
zu = m ' . 7 ++ ee A ANTE 
A ee WER er erringen Ves ee NS, Dent i hi Fe ors ee Stites PIU 


= {TORE FAR arg) is A 
Po DE le: Met Nano my ЗН 
SNL oe eae A 


ix 
Per liege AAA 


a a ave 
4 Mecque Berre hrs 


ot AISA, rado» RTL 
LE ee 
= "Maires de 


MALACOLOGIA, 2008, 50(1-2): 1-12 


TAXONOMY, COMPARATIVE MORPHOLOGY, AND GEOGRAPHICAL 
DISTRIBUTION OF THE NEOTROPICAL GENUS HYPSELARTEMON WENZ, 1947 
(GASTROPODA: PULMONATA: STREPTAXIDAE) 


André Е. Barbosa", Norma С. Salgado? & Arnaldo С. dos Santos Coelho? 


ABSTRACT 


The diagnosis and geographical distribution of the genus Hypselartemon Wenz, 1947, 
which was originally proposed as a subgenus of Rectartemon Baker, 1925, were re-exam- 
ined based on preserved material, including type specimens, collected since the nine- 
teenth century and deposited in scientific institutions. The comparative morphology of shell, 
radula, and soft body parts, when available, of Hypselartemon alveus (Dunker, 1845), H. 
contusulus (Férussac, 1827), H. deshayesianus (Crosse, 1863), and H. paivanus (Pfeiffer, 
1867) were analyzed in order to redescribe and to highlight characters that can be used 
for species identification. Examination of the reproductive organs revealed significant dif- 
ferences among the species, all of which are endemic to the state of Rio de Janeiro, Brazil, 
and do not occur in Colombia as originally suggested. These results seem useful in exam- 
ining the taxonomy and systematics of the Neotropical Streptaxidae. 

Key words: Streptaxidae, Hypselartemon, taxonomy, morphology, neotropics. 


INTRODUCTION 


The family Streptaxidae includes carnivorous 
land snails distributed throughout the tropics 
and subtropics (Bruggen, 1967). Most of the 
Brazilian Streptaxidae were described in the 
19th and first half of the 20th centuries by for- 
eign malacologists, such as Férussac (1827), 
Pfeiffer (1849, 1850a, 1867), Pilsbry (1897, 
1930), and Baker (1914). The descriptions 
were often based only on shell characters, 
usually without illustrations. The synopses of 
Gude (1902), Kobelt (1910), and Richardson 
(1988) are still the most comprehensive pa- 
pers on the South American Streptaxidae spe- 
cies and provide references indispensable for 
work on the family. 

Due the scarce understanding of the internal 
anatomy of Streptaxidae, most genera that 
occur in Brazil still have taxonomic problems 
and remain diagnosed only by shell charac- 
ters. Hypselartemon Wenz, 1947, was origi- 
nally described as a subgenus of Rectartemon 
Baker, 1925, with Rectartemon (Hypse- 
lartemon) alveus (Dunker, 1845) as the only 
species explicitly included. Wenz (1947) sug- 


gested that Hypselartemon might include more 
species, but without listing them, with a distri- 
bution restricted to Brazil and Colombia. 

Despite being characterized by Zilch (1960) 
and treated as a valid subgenus by Vaught 
(1989), Hypselartemon as well as Rectartemon 
were considered as junior synonyms of Artemon 
Beck, 1837, by Richardson (1988). Indeed, 
there has long been disagreement regarding 
the taxonomic validity of Streptaxis Gray, 1837, 
Artemon, and Rectartemon, as discussed by 
Deshayes (1851), Baker (1925), Thiele (1927), 
Pilsbry (1930), and Solem (1956). 

Barbosa et al. (2002) redescribed Helix 
contusula Férussac, 1827, with a new generic 
arrangement in Hypselartemon, a status sug- 
gested by Salgado & Coelho (2003) to include 
the following species with an ovate shell and a 
high spire: Hypselartemon alveus (Dunker, 
1845), H. deshayesianus (Crosse, 1863), and 
H. paivanus (Pfeiffer, 1867). 

Here, we recharacterize the genus Hypse- 
lartemon and all species using conchological 
data and the morphology of soft body parts, 
when available, in order to improve the taxon- 
omy of the Brazilian Streptaxidae. 


‘Departamento de Malacologia, Instituto Oswaldo Cruz, Av. Brasil 4365, 21040-900 Rio de Janeiro, Brazil 
?Malacologia, Departamento de Invertebrados, Museu Nacional - Universidade Federal do Rio de Janeiro, Quinta da Boa 


Vista, 20940-040, Rio de Janeiro, Brazil 
“Corresponding author: andrefbarbosa@gmail.com 


METHODS 
Institutional Abbreviations 


Specimens examined are found in the mol- 
lusc collections of the Academy of Natural 
Sciences of Philadelphia, Pennsylvania, 
U.S.A. (ANSP), Muséum National d’Histoire 
Naturelle, Paris, France (MNHN), Museu 
Nacional, Universidade Federal do Rio de 
Janeiro, Brazil (MNRJ), and Senckenberg- 
Museum, Frankfurt am Main, Germany (SMF). 


Morphological Examination 


Terminology for shell and soft body parts 
were adopted from Rezende et al. (1962), 
Berry (1965), and Picoral & Thomé (1998). For 
adult shells, the following characters were 
measured using a Cannon Caliper Rule: H: 
total shell height; Hs: spire height; DM: major 
shell diameter; dm: minor shell diameter; Du: 
major umbilicus diameter; Ha: aperture height; 
Da: major aperture diameter. Modal values of 
dimension were used in the diagnosis. The 
number of whorls was determined according 
to Diver (1931). 

Shell illustrations were obtained from differ- 
ent sources — a digital camera (MNHN types; 
Figs. 7, 10, 13), a camera coupled to a Zeiss 
SV11 stereomicroscope (Fig. 13), scanned 
from original illustrations (Fig. 4) or micro- 
graphs taken with a LEO 1450 VP electron 
microscope (Figs. 5, 6, 8, 9, 11, 12). 

Living specimens were drowned in water and 
preserved in 70% ethanol. Soft parts were iso- 
lated after crushing the shells, and were dis- 
sected under a light microscope (Zeiss Stemi 
SV11) that was coupled to a camera lucida. 


RESULTS 


The most important conchological criteria 
used to diagnose the species were the spire 
height and its relation to total height of shell 
and the body whorl deflection. A detailed char- 
acterization of H. contusula, including its soft 
parts, was previously published by Barbosa 
et al. (2002). 


Hypselartemon Wenz, 1947 


[Greek: hypselos, high; artemon, pulley] 

Rectartemon (Hypselartemon) Wenz, 1947: 36 
— Zilch, 1960: 558; Vaught, 1989: 91. 

Hypselartemon Wenz, 1947 — Barbosa et al., 
2002: 2; Salgado & Coelho, 2003: 170. 


BARBOSA ET AL. 


Type Species 
Streptaxis alveus Dunker, 1845, by original 
designation; Brasilia; Neufreiburg, Prov. Rio 
Janeiro. 


Distribution Originally Suggested 
“Brasilien, Columbia’. 


Distrubution Here Suggested 
Brazil, Rio de Janeiro state. 


Included Species 

Hypselartemon alveus (Dunker, 1845), H. con- 
tusulus (Ferussac, 1827), H. deshayesianus 
(Crosse, 1863), and H. paivanus (Pfeiffer, 1867). 


Original Diagnosis 

“Gehäuse kreiselförmig, etwa so hoch wie 
breit, mit stärker (als bei Rectartemon s. str.) 
erhobenem Gewinde und 8-10 Umgängen. D 
= 10-28 mm”. 


Diagnosis 

Shell whitish in color, small, ovate to 
cupuliform, robust, with prominent spire, almost 
as high as wide, 6-10 convex whorls. Body 
whorl flattened to convex, slightly deviating or 
not from columellar axis. Peristome strongly 
reflected. Н = 5.0-13.0 mm; DM = 5.0-9.5 mm. 


Hypselartemon alveus (Dunker, 1845) 
(Figs. 1-3) 


Streptaxis alveus Dunker, 1845: 7, pl. 6, fig. 
14 — Pfeiffer, 1848a: 4; 1850b: 15, pl. 101, 
figs. 33-35; Hupé, 1857: 20; Gude, 1902: 
207-208; Kobelt, 1905: 29-30. 

S. [treptaxis] alveus Dunk. [section Artemon”] 
— Pfeiffer & Clessin, 1881: 15. 

Streptaxis (Eustreptaxis) alveus Dunker — 
Tryon, 1885: 69, pl. 27, fig. 11; Kobelt, 1910: 
144. 

S.[treptaxis] (А. [rtemon]) alveus Dunker — 
Thiele, 1931: 729. 

Alcidia alveus (Philippi) — Bourguignat, 1889: 7. 

Artemon alveus (Dunker, 1845) — Lange-de- 
Morretes, 1949: 166; Richardson, 1988: 174. 

Rectartemon (Hypselartemon) alveus 
(Dunker) — Wenz, 1947: 36; Zilch, 1960: 558, 
МО. 1953; 196% 81, 

Hypselartemon alveus (Dunker, 1845) — 
Barbosa et al., 2002: 2-3; Salgado & Coelho, 
2005 170: 


Type Locality 
“Neufreiburg, Prov. Rio Janeiro” (Dunker, 
1845). 


TAXONOMIC REVISION OF THE GENUS HYPSELARTEMON 


FIGS. 1-13. Hypselartemon spp. shell. FIGS. 1-3: H. alveus, ANSP 23728; frontal, basal (Н = 12.2 
mm; DM = 9.4 mm) and juvenile (H = 8.0 mm; DM = 7.6 mm); FIG. 4: Helix contermina; FIGS. 5-7: H. 
contusulus, MNRJ 8436 (5-6; H = 6.4 mm; DM = 5.9 mm) and MNHN (7 — Lectotype; H = 6 mm; DM 
= 5.1 mm); FIGS. 8-10: H. deshayesianus, MNRJ HSL3395 (8-9; H = 5.5 mm; DM = 6 mm) and 
MNHN (10 — Syntype; H = 5 mm; DM = 5.5 mm); FIGS. 11-13: H. paivanus, MNRJ 8350 with umbilicus 
detail (11-12; H=5.7 тт; DM = 6.8 mm) and MNHN (13 — Syntype; H = 5.0 mm; DM = 6.0 mm). Photo 
1 by A. F. Barbosa; photos 2-3 by P. M. S. Costa; photo 4 after Tryon (1885: pl. 12, fig. 19); photos 5- 
6; 8-9; 11-12, electron micrographs; photos 7; 10 and 13 by P. Maestrati. Scale bars: 1 mm. 


é. BARBOSA ET AL. 


Distribution 
Brazil, Rio de Janeiro state, Nova Friburgo. 


Diagnosis 

Shell with very high spire (Hs: 7.5 mm), body 
whorl short, flat at its base, without deviation 
from columellar axis. 


Description 

Shell (Figs. 1-3): Triangular, taller than wider, 
robust, whitish, with 872-10 slightly convex 
whorls. Protoconch with 11/3 whorls, usually 
smooth, sometimes with some granulation or 
lightly striated. Spire very high. Suture shal- 
low. Next to half of second whorl there is a 
deeply delimited border; from there whorls 
become wider, less convex, almost flat, with 
deeply marked striae. Body whorl short, flat at 
its base, about 1/5 of total shell height, without 
deviation from columellar axis, striated until 
umbilicus. Aperture luniform, descendant, long, 
narrow; peristome reflected, expanded from its 
base to columellar lip, where it enlarges and 
covers 1/3 of narrow umbilicus; parietal lip flat, 
external lip with elliptical base. 

Dimensions (тт): H: 10-12.4; Hs: 7.3-7.7; 
DM: 9.0-9,4; dm: 8.5-9.0; Aperture: Ha: 4.0- 
4.3; Da: 3.6-4.0. Umbilicus: Du: 0.6. 


Remarks 

Helix contermina Reeve, 1854 (pl. 191, sp. 
1342) (Fig. 4), originally described with 6-7 
whorls, pupiform, conic-globose, convex at its 
base and from an unknown locality, was con- 
sidered as a junior synonym of H. alveus by 
Pfeiffer (1859), Pfeiffer & Clessin (1881), 
Tryon (1885), Gude (1902), Kobelt (1905, 
1910), and Richardson (1988). We compared 
the original diagnosis and shell illustration of 
H. contermina and its later copy by Tryon 
(1885) with shells and illustrations of H. 
alveus. These two species differ in that H. 
contermina has almost half the number of 
whorls and shell height, as well as a convex 
body whorl base, contrasting with the flat and 
plane body whorl base of H. alveus. Even 
young shells of H. alveus, the size of which 
are close to that of H. contermina, have a 
notable flat body whorl base just like adult 
shells. We were not able to locate the type 
material of H. contermina in any ofthe many 
museums consulted. Despite being consid- 
ered a junior synonym of H. alveus by many 
authors, we consider the available data doubt- 
ful and insufficient to confirm its taxonomic 
position, until type material of H. contermina 
can be found and properly examined. 


Juvenile shells of H. alveus were also exam- 
ined and have similar dimensions as H. de- 
shayesianus and H. contusulus. They differ in 
that H. alveus has a very short and flat body 
whorl, whereas the shells of H. deshayesianus 
and H. contusulus are more convex. This infor- 
mation might be useful in the taxonomic iden- 
tifcation of Hypselartemon shells in collections 
without precise indication of the collect locality. 

Neither living specimens nor empty shells 
of H. alveus have been found in recent collec- 
tions in the vicinity of its type locality. 


Material Examined 

Brazil (without more precise locality), ANSP 
4353, 1 shell, A. D. Brown coll.; ANSP 23727, 1 
shell, Swift coll.; Anthony coll.; ANSP 23728, 4 
shells, J. S. Phillips coll. SMF 136864, 1 shell, 
Kobelt colin. ["typoide”? — Orig. fig., Kobelt (1905) 
and Zilch (1960), according to Zilch (1961)]. 


Hypselartemon contusulus (Férussac, 1827) 
(Figs. 5-7) 


Helix (Helicogena) contusula Férussac, 1827: 
302 — Rang, 1831: 9; Chevalier, 1966: 1009. 

A.[rtemon] contusulus (Еег.) — Beck, 1837: 48. 

Hypselartemon contusulus (Férussac, 1827) 
— Barbosa et al., 2002: 1-10. 


Type Locality 
“Rio-Janeiro” (Férussac, 1827). 


Distribution 

Rio de Janeiro State, (...) “dans les bois et 
sous les baies” (Rang, 1831); Angra dos Reis, 
Ilha Grande (Barbosa et al., 2002). 


Diagnosis 

Shell with high spire (Hs: 3.5 mm) and con- 
vex body whorl that slightly deviates from col- 
umellar axis. Cylindrical penial complex 
slender, with or without spines. Free oviduct 
cylindrical, rectilineal: vas deferens wide at its 
insertion with prostate, narrow where it 
emerges from penial muscular sheath. Penis 
with long, muscular sheath, thick, strong. 


Description 

See Barbosa et al. (2002). 

Dimensions (тт): H: 5.0-6.8; Hs: 3.3-3.6; 
DM: 5.0-6.4; dm: 4.8-5.3. Aperture: Ha: 2.2- 
3.0; Ва: 2.4-2.8. Umbilicus: Du: 0.6-0.8. 


Remarks 
The species was first collected by Sander 
Rang, an officer of the French Royal Navy, 


TAXONOMIC REVISION OF THE GENUS HYPSELARTEMON 5 


during the Great Indias Expedition, and sent 
to Ferussac, who described it without any il- 
lustration in an obscure journal (Férussac, 
1827). Chevalier (1966) noted that H. 
contusula was not mentioned in Férrussac's 
Histoire Naturelle des Mollusques (Ferussac 
& Deshayes, 1819-1851). As a result, the spe- 
cies remained unrecorded for over a century 
in the malacological literature. It was re- 
described by Barbosa et al. (2002) based on 
syntypes in the MNHN and material recently 
collected in Ilha Grande, Angra dos Reis, State 
of Rio de Janeiro, Brazil. 

Although not mentioned in the recharacter- 
ization by Barbosa et al. (2002), the presence 
of external spines in the middle portion of phal- 
lus of H. contusulus was observed in one dis- 
sected specimen. These structures are 
common in Streptaxidae, as well as in other 
Stylommatophora, and were discussed by 
Berry (1965), Gerlach (1995), and Picoral & 
Thome (1998). 


Material Examined 

Lectotype (designated by Barbosa et al., 
2002): Brazil, Rio de Janeiro, MNHN 1 shell, 
Rang coll.; Paralectotypes: same designation 
and data as lectotype, MNHN 2 shells (1 
young); Angra dos Reis, Ilha Grande (23°11’S, 
44°12’W), Parnaioca beach footpath, on or 
buried under the soil and among dead leaves 
and decaying material, MNRJ 8436, 1 shell, 
N. C. Salgado & S. B. Santos colls., 15/VIII/ 
1996; MNRJ 8440, 3 specimens, N. C. 
Salgado & $. В. Santos colls., 27/IV/1996; 
MNRJ 8442, 6 specimens (1 young), S. B. 
Santos & V. Queiroz colls., 08/V/1997; MNRJ 
8438, 5 specimens (1 young), A. F. Barbosa, 
S.B. Santos & P. M. Coelho colls., 16/1/1998; 
MNRJ 8439, 2 specimens, A. F. Barbosa, S. 
В. Santos & К. $. Massa colls, 17/1/1998; 
ММК} 8437, 1 specimen, А. Е. Barbosa coll., 
16/1/1999; MNRJ 7727, 5 specimens (3 
young), S. В. Santos coll., V/1997; MNRJ 
7717, 2 shells, S. B. Santos coll., 30/V/1997; 
MNRJ 8441 soft parts + 1 roof of pallial cavity 
+ 2 systems (reproductor and digestive). Ilha 
Grande, Parnaioca beach footpath, Toca das 
Cinzas: MNRJ 7732, 2 shells, S. B. Santos & 
V. Queiroz colls., 13/VIII/1996; ММК} 8443, 3 
specimens, A. F. Barbosa, S. B. Santos & P. 
M. Coelho colls., 17/1/1998. Ilha Grande, 
Cachadaco, ММК} 7702, 9 shells (3 frag- 
mented), S. B. Santos e V. Queiroz colls., 
30/V/1997. 


Hypselartemon deshayesianus (Crosse, 1863) 
(Figs. 8-10) 


Streptaxis deshayesianus Crosse, 1863: 388 
— Crosse, 1867: 202, pl. 5, fig. 3; Gude, 1902: 
208; 1903: 325; Hidalgo, 1870: 39; 1872: 45- 
46, pl. 3, figs. 5, 6; Kobelt, 1905: 30, pl. 46, 
figs. 45; Richardson, 1988: 253-254. 

Streptaxis deshayesianus Crosse — Tryon, 
1885: 69, pl. 14, fig. 97 in “section Eustrep- 
taxis’; “subsection Edentulae”. 

Streptaxis (Eustreptaxis) deshayesianus 
Crosse — Kobelt, 1910: 145. 

Artemon deshayesianus (Crosse, 1863) — 
Lange-de-Morretes, 1949: 166. 

Hypselartemon deshayesianus (Crosse, 1863) 
— Barbosa et al., 2002: 3; Salgado & Coelho 
2003170. 


Type Locality 
“Habitat ?” (Crosse, 1863). 


Distribution 

Rio de Janeiro, “Botafogo, dans le 
Corcobado, a Rio Janeiro (Paz et Martinez)” 
(Hidalgo, 1870) and Barra da Tijuca. 


Diagnosis 

Shell small, with short spire (Hs: 2.8 mm), 
and convex body whorl that slightly deviates 
from the columellar axis. 


Description 

Shell (Figs. 8-10): Cupuliform, as high as 
wide, thin, whitish, translucent, with 672-7 very 
convex whorls. Spire short. Protoconch with- 
out defined limit, first whorls smooth; next to 
1/3 Of second whorl there is a deep delimited 
border; from there, whorls become wider but 
remain very convex, with regular, strongly 
marked axial striation to shell base. Suture 
very deep. Body whorl convex at its base, 
short, about % of total shell height, briefly de- 
viating from columellar axis, striated until um- 
bilicus, which is wide, deep. Aperture oval, 
descendent, not very long; peristome round, 
reflected, expanded from base to columellar 
lip; parietal lip convex, external lip oval. 

Dimensions (mm): Н: 5.7-6.0; Hs: 2.7-2.9; 
ОМ: 5.8-6.0; ат: 5.0-5.4. Aperture: Ha: 2.1- 
2.5; Da: 2.5-2.6. Umbilicus: Du: 0.5-0.7. 


Remarks 
Shells of this species were obtained during 
the Spanish scientific expedition to Meridional 


6 BARBOSA ET AL. 


America, from 1862 to 1865, headed by Don 
Patricio Maria Paz y Membiela, with the ob- 
jective increasing the collections of the Madrid 
museum (Hidalgo, 1872). Sent to Crosse, the 
species was described in 1863 but not illus- 
trated until 1867. The locality of this material 
was unknown to Crosse. Hidalgo (1870, 1872) 
indicated the locality of the shells according 
to records of Paz and Martinez, members of 
the Spanish expedition. According to Paz 
(Hidalgo, 1870), within two hours, more than 
100 specimens of the following species were 
easily collected: “Streptaxis crossei, S. 
paivanus and $. deshayesianus”. The natural 
habitats of H. deshayesianus have been 
strongly impacted by human activity, being 
located in the second largest urban area in 
Brazil. Neither living specimens nor empty 
shells of H. deshayesianus have been found 
in recent collections in the city of Rio de 
Janeiro. 

As early as 1872, Hidalgo observed that H. 
deshayesianus was very similar to H. alveus. 


Material Examined 

Brazil, Rio de Janeiro, MNHN (syntypes, 
according to label information) 2 shells, M. 
Hidalgo coll.; Barra da Tijuca, ММК HSL3395, 
4 shells, Н. $. Lopes coll., VI11/1952. 


Hypselartemon paivanus (Pfeiffer, 1867) 
(Figs. 11-15) 


Streptaxis paivana Pfeiffer, 1867: 43, pl. 1, 
fig: 2: 

Streptaxis paivanus Pfeiffer — Hidalgo, 1870: 
39; 1872: 44-45, pl. 3, figs. 3, 4; Gude, 
1902: 230; 1903: 326. 

Streptaxis paivanus Pfr. — Pfeiffer & Clessin, 
1881. 15; 180n;, 188562, pl, 12, figs: 7, 
8, in “section Artemon” 

Streptaxis (Streptartemon) paivanus L. Pfr. 
— Kobelt, 1905: 37-38, pl. 46, figs. 6, 7; 
1910: 145. 

Alcidia paivana (Pfeiffer) — Bourguignat, 
1889: 47. 

Artemon paivanus (Pfeiffer, 1867) — Lange- 
de-Morretes, 1949: 166. 

Streptartemon paivanus (Pfeiffer, 1867) — 
Richardson, 1988: 250. 

Hypselartemon paivanus (Pfeiffer, 1867) — 
Barbosa et al., 2002: 3; Salgado & Coelho, 
2003:.1.70, 


Type Locality 
“Hab. in Brasilia loco Macahe dicto” (Pfeiffer, 
1867). 


Distribution 
Brazil, Rio de Janeiro State: Macaé, Buzios, 
Cabo Frio, Arraial do Cabo and Araruama. 


Diagnosis 

Shell with short spire (Hs: 3.3 mm) and wide 
diameter. Short body whorl slightly convex at 
its base, not deviating from the columellar axis. 
Umbilicus very deep, wide. Penial complex 
wide, bearing corneous spines; free oviduct 
wide, curved; vas deferens with narrow diam- 
eter along its entire length. Sheath of penis 
thin, membranous, short. 


0,5mm 


FIG. 14. Hypselartemon paivanus, roof of pallial 
cavity. АМ — anus; AU — auricle; IN - intestine; KI 
— kidney; PC — pericardium; PU — primary ure- 
ter; PV — pulmonary vein; RE — rectum; SU — 
secondary ureter; VE — ventricle. 


TAXONOMIC REVISION OF THE GENUS HYPSELARTEMON Z 


Description 

Shell (Figs. 11-13): Cupuliform, wider than 
tall, whitish, translucent, thin, with 7-8 Y con- 
vex whorls. Protoconch without defined limit. 
Spire very short. Deep suture. First whorls 
smooth; next to 5/5 of the second whorl there 
is a deep delimited border; from there whorls 
become more convex and wider, with strongly 
marked regular axial striae that continue to 
shell base. Body whorl slightly convex, short, 
about 1/z of the total shell height, not deviating 
from columellar axis. Aperture circular, descen- 
dent, not long; parietal lip convex; peristome 
round, not expanded, reflected from base to 
columellar lip. Umbilicus very wide, deep. 


Dimensions (mm): H: 5.4-6.4; Hs: 3.2-3.4; 
DM: 6.2-7.8; dm: 6.3-6.7. Aperture: Ha: 2.5— 
3.0; Da: 3.2-3.3. Umbilicus: Du: 1.0-1.2. 

Radula: Long and slender ribbon, with about 
45 sharply pointed unicuspid teeth in trans- 
verse row (22-1-22), size increasing from 
marginal to lateral. Central teeth reduced, slen- 
der, curved. 

Roof of Pallial Cavity (Fig. 14): Long, slen- 
der, narrow, strongly wrinkled, macroscopic 
venation not conspicuous, except for the wide 
pulmonary vein, which leaves the pericardium 
and approaches the mantle edge. Heart pear- 
shaped, auricle and ventricle well defined. Kid- 
ney distally globose with constriction in 


Po 


FIG. 15. Hypselartemon paivanus, reproductive system anatomy. AG — albumen 
gland; AT — atrium; BC — bursa copulatrix; BD — bursa copulatrix duct; CD — 
collecting duct; DD — vas deferens; DG - digestive gland; EP - epiphallus; FC — 
fertilization complex; FO - follicles; HD — hermaphroditict duct; OD - free ovi- 
duct; PH — phallus; PR — prostate gland; PS — penial sheath; RM — penial retrac- 
tor muscle; SV — seminal vesicle; UT — uterus; VA — vagina. 


8 BARBOSA ET AL. 


proximal part, where the sigmoid-shaped pri- 
mary ureter exits, following the margin of the 
kidney until the origin of the secondary ureter, 
which follows close to the rectum until mantle 
edge. 

Reproductive Organs (Fig. 15): Ovotestis 
embedded in the digestive gland, consisting 
of follicle groups that open into the collecting 
duct. Hermaphroditict duct long, narrow. Semi- 
nal vesicle emerging from median portion of 
hermaphroditict duct, consisting of a sinuous 
blind tube similar in width to the 
hermaphroditict duct. Fertilization complex 
embedded in albumen gland, externally con- 
stituted of a hermaphroditict duct fold and al- 
bumen gland duct. Spermoviduct with wrinkled 
uterus and prostate gland. Free oviduct large, 
curved, C-shaped, narrower where it inserts 
in spermoviduct. Bursa copulatrix duct 
emerges near middle of the free oviduct, far 
from the point where vas deferens emerges 
from prostate. Phallus narrow next to the 
atrium and wider in its middle part up to the 
epiphallus, where it constricts at the top. It is 
armed with small external corneous spines, 
notably on its middle part. Penial retractor 
muscle slender, narrow, similar in width to vas 
deferens, very long, approximately as long as 
phallus and epiphallus together. Narrow vas 
deferens emerges from prostate, follows free 
oviduct and dive under penial sheath until its 
terminal portion, where it folds and emerges 
again, coupled to phallus. Then, it follows all 
the phallus extension in a sinuous or rectilineal 
trajectory, inserting at epiphallus constriction, 
near insertion of muscular retractor penis. 
Penial sheath thin, membranous, of 1/5 of to- 
tal length of penial complex. Atrium globose, 
vagina of same width as first third of penis. 


Remarks 

Specimens of H. paivanus are found in ar- 
eas near beaches, with sandy soil and typical 
“restinga” vegetation (costal dune forest). Their 
shells can be found in relative abundance, but 
living specimens are rare. In its natural habi- 
tat, other terrestrial molluscs were collected 
and could form part of the diet of H. paivanus, 
include species of Helicinidae, Subulinidae 
and juvenile Bulimulidae and Strophocheilidae. 


Material Examined 

Brazil, Rio de Janeiro, MNHN (syntypes, 
according to label information) 3 shells (1 from 
Cabo Frio), Journ. Conchy. coll., M. Hidalgo 
coll.; Arraial do Cabo, MNRJ 8349, 7 speci- 
mens + 1 cephalopedious mass + 1 radula, B. 


5. Dunley coll., 9/11/2000; Prainha, MNRJ 
8349, 4 shells + 3 reproductive systems + 2 
roof of palial cavity + 1 nervous system + 1 
bucal mass, С. J. Е. Costa col., 03/X1/2001; 
Praia do Forno, MNRJ 7542, 2 shells, P. 
Jurberg col., 14/IV/1963; MNRJ 8350, 3 shells, 
A. F. Barbosa & V. Bessa colls., 21/VII/2000; 
MNRJ 9750, 16 shells, J. C. Monteiro col., 
18/1/2001; Buzios, Praia da Tartaruga, MNRJ 
8351, 3 shells, A. F. Barbosa & V. Bessa colls.; 
22/V11/2000. 


DISCUSSION 


Hypselartemon was considered a subgenus 
of Rectartemon by Zilch (1960) and Vaught 
(1989) and a synonym of Artemon by 
Richardson (1988). Following Barbosa et al. 
(2002) and Salgado & Coelho (2003), we con- 
sider Hypselartemon to have generic status 
based on its shared morphological characters. 

Shells data including dimensions, shape, 
spire height and peristome reflection obtained 
by comparative analysis in four species were 
in accordance with the Hypselartemon origi- 
nal diagnosis (Wenz, 1947), reviewed by Zilch 
(1960) and Barbosa et al. (2002). Earlier, 
Hidalgo (1872) already noticed a strong mor- 
phological resemblance of H. a/veus and Н. 
deshayesianus. The similar appearance of 
both species led Bourguignat (1889) to group 
H. alveus and H. paivanus in a separete ge- 
nus Alcidia Bourguignat, 1889. 

In comparison with other Brazilian 
Streptaxidae genera, Hypselartemon can be 
considered a well-defined group — its shell is 
not strongly detorted, as in Streptaxis or 
Streptartemon Kobelt, 1905; it is smaller in 
diameter and proportionally higher than 
Rectartemon or Artemon and Martinela 
Jousseaume, 1887; and it has a circular aper- 
ture differing from the monotypic Sairostoma 
Haas, 1938. 

The dimensions originally proposed for 
Hypselartemon shells (‘D = 10-28 mm”) were 
not completely congruent with the dimensions 
found in the species that we studied. Hypse- 
lartemon alveus, which is the type species of 
the genus and the only species recognized by 
Wenz (1947), has a diameter of 9.4 mm, which 
is close to the lower limit suggested in the 
Hypselartemon original diagnosis. The upper 
limit of 28 mm mentioned by Wenz (1947) is 
not found in any of the species included in the 
genus. Wenz (1947) probably went too far in 
estimating those dimensions, once a shell with 


TAXONOMIC REVISION OF THE GENUS HYPSELARTEMON 


a diameter of 28 mm and and a tall spire, as 
diagnostic for Hypselartemon, has never been 
described as Streptaxidae species in South 
America. 

The lower limit of the number of whorls of 
Hypselartemon shells originally indicated (8 
whorls) also did not include the number of 
whorls in two of the four species examined: 
H. contusulus (6-77 whorls) and H. 
deshayesianus (672-7 whorls). 

The shell morphology of H. contusulus is very 
similar to that of H. deshayesianus considering 
the number of whorls and shell height, differing 
in that the former has a more convex body whorl 
that deviates less from the columellar axis, and 
a smaller umbilicus. Hypselartemon alveus and 
H. paivanus have a higher number of whorls 
(972 and 8% respectively), the body whorl does 
not deviate from the columellar axis, and the 
spire is flattened at its base. Hypselartemon 
alveus is the tallest species (H: up to 12.7 mm). 
Hypselartemon paivanus has the largest shell 
width compared to shell height (DM: 7.2 mm) 
(Barbosa et al., 2002). 

Examination of the soft parts, mainly of the 
reproductive system, revealed significant dif- 
ferences between H. paivanus and H. 
contusulus (Table 1). The group numbers and 
arrangement of the follicles in ovotestis, the 
shape and size of the seminal vesicle, and the 
length and absence of ramifications of the 
penial retractor muscle are similar in these two 
species. 

The roof of pallial cavity of H. paivanus is 
similar to Pilsbry’s (1907) illustration for 
Streptartemon deformis (Férussac, 1821). It 
differs from H. contusulus in that the latter has 
a more conspicuous macroscopic venation. 
Both are long, slender, narrow, and wrinkled, 
as is common for Streptaxidae. 


co 


Structure and form of the radula of H. 
paivanus are similar to that of H. contusulus, 
as examined by Barbosa et al. (2002), and dif- 
fers basically in the number of teeth per trans- 
verse row. Solem (1974) provided an 
examinaton on the diversity of radular modifi- 
cations among carnivorous land snails and its 
implications on feeding habits. The relevance 
of Brazilian Streptaxidae radula for species di- 
agnosis is not clear yet. 

In his introduction to Streptaxidae, Bruggen 
(1967) noted that the anatomical data about 
the family are “few and far between”. The lack 
of soft part informations is still a problem for 
worldwide malacologists working on 
Streptaxidae, as recently argued by Clements 
(2006). The bulk of Streptaxidae specimens 
deposited in museums collections are only 
empty shells. Generic definitions and diagnos- 
tics characters for Brazilian Streptaxidae re- 
mains based on shell data. As common for 
Stylommatophora, characters of genitalia are 
most informative for its taxonomy and phylog- 
eny. However, due to the lack of soft parts of 
two Hypselartemon species examined, we 
were not able to use soft parts characters in 
the generic diagnosis, until more work on com- 
parative morphology in the family can be prop- 
erly done. 

The original geographical distribution sug- 
gested for Hypselartemon (Brazil and Colom- 
bia) could not be confirmed according to the 
species included until now in this genus. The 
search for Hypselartemon speciemens in mol- 
lusc collections in Colombia (Universidad 
Nacional de Colombia, Instituto de Ciencias 
Naturales and Instituto Alexander von 
Humboldt) was unsuccessful. Published 
records and collection data indicate that spe- 
cies of Hypselartemon are endemic to the state 


TABLE 1: Summary of the reproductive system differences found in two species examined. 


Characters 


Hypselartemon paivanus 


Hypselartemon contusulus 


Free oviduct 


Vas deferens narrow in its entire length 


Penial complex 


Penial sheath short, thin, membranous 


Bursa copulatrix duct emerges far from ovispermoduct 


large, strongly curved, C-shaped 


narrow, cylindrical, rectilinear, long 


emerges with larger diameter from 
penial muscular sheath 


enlarged with epiphallus constricted on cylindrical, long, narrowed at 
top, always bearing external spines 


epiphallus, bearing external spines 
or not 


long, thick, muscular, strong 
emerges next to ovispermoduct 


10 BARBOSA ET AL. 


of Rio de Janeiro because from the 19" cen- 
tury onwards they have been collected only 
there. These data can be reinforced by exten- 
sive field work of our team and collaborators 
all around Rio de Janeiro and other states in 
Brazil. The increasing human impact in areas 
where species of Hypselartemon are found 
may represent a threat to their conservation 
because of extensive habitat loss and frag- 
mentation of suitable natural areas. 


ACKNOWLEDGEMENTS 


We thank Dr. Charlene Fricker from the 
Academy of Natural Sciences of Philadelphia, 
Pennsylvania, USA, and Philippe Maestrati, 
Museum National d’Histoire Naturelle, Paris, 
France, for providing access and photographs 
of vouchers and type specimens; the technolo- 
gist J. Eduardo Prado from the Malacology 
Laboratory of the Instituto Oswaldo Cruz, Rio 
de Janeiro, for preparing the genitalia anatomi- 
cal drawing. Jonas D. Brito and MSc Victor 
Wagner S. Lopes, of the Universidade do 
Estado do Rio de Janeiro, prepared the elec- 
tron micrographs; and Dr. Julio С. Monteiro 
and Claudio J. F. Costa, from the Museu 
Nacional, Universidade Federal do Rio de 
Janeiro, for collecting important samples of liv- 
ing H. paivanus used for anatomical studies. 
W. Lobato Paraense, from Instituto Oswaldo 
Cruz, and Malacologia’s referees and editor 
kindly reviewed the manuscript. The senior 
author is supported by a SVS/Fiocruz fellow- 
ship. 


LITERATURE CITED 


BAKER, F. C., 1914, The land and fresh-water 
mollusks of the Standford Expedition to Brazil. 
Proceedings of the Academy of Natural Sci- 
ences of Philadelphia, 65: 618-672. 

BAKER, H. B., 1925, The Mollusca collected by 
the University of Michigan-Williamson Expedi- 
tion in Venezuela. Occasional Papers of the 
Museum of Zoology, 156: 1-57. 

BARBOSA, А. F., М. С. SALGADO & А. С. $. 
COELHO, 2002, Taxonomic status and rede- 
scription of Helix contusula Férussac, 1827 as 
Brazilian Streptaxidae species (Mollusca, Gas- 
tropoda, Pulmonata). Boletim do Museu 
Nacional, n. s., Zoologia, 482: 1-10. 

BECK, H. H., 1837-1838, Index molluscorum 
praesentis aevi musei principis augustissimi 
Christiani Frederici, fasciculus primus. Mol- 
lusca gastraeopoda pulmonata. Hafniae, pp. 
1-100 (1837), 101-124 (1838). 


BERRY, A. J., 1965, The genital systems of Dis- 
cartemon stenostomus Van Benthen Jutting 
and Huttonella bicolor (Hutton) (Pulmonata, 
Streptaxidae) from Malaya. Proceedings of the 
Malacological Society of London, 36(4): 221- 
228. 

BOURGUIGNAT, J. R., 1889, Mollusques de Г 
Afrique équatoriale de Moguedouchou à 
Bagamoyo et de Bagamoyo au Tanganika. 
Paris, 229 pp. 

BRUGGEN, A. C. VAN, 1967, An introduction to 
the pulmonate family Streptaxidae. Journal of 
Conchology, 26(3): 181-188. 

CHEVALIER, H., 1966, Catalogue des collections 
du Muséum correspondant a L’ “Histoire 
Naturelle des Mollusques” de Férussac (4° 
partie). Bulletin du Muséum National d’Histoire 
Naturelle, 37(6): 1001-1012. 

CLEMENTS, R., 2006, Oophana tiomanensis, a 
new Streptaxidae (Gastropoda: Pulmonata: 
Streptaxidae) from Pulau Tioman, Peninsular 
Malaysia. The Raffles Bulletin of Zoology, 
54(1): 125-127. 

CROSSE, H., 1863, Diagnoses d’ espéces 
nouvelles. Journal de Conchyliologie, 11(4): 
388-389. 

CROSSE, H., 1867, Descriptions d’espéces 
nouvelles. Journal de Conchyliologie, 15(2): 
195-203. Е 

DESHAYES, С. Р.; see FERUSSAC & DE- 
SHAYES. 

DIVER, C., 1931, A method of determining the 
number of whorls of the shell and its applica- 
tion to Cepaea hortensis Mull. Proceedings of 
the Malacological Society of London, 19(5): 
234-239. 

DUNKER, W., 1845, in: R.A. PHILIPPI, 1845-1851, 
Abbildungen und Beschreibungen neuer oder 
wenig gekannter Conchylien, unter Mithülf 
mehrerer Deutscher Conchyliologen, 
herausgegeben von R. A. Philippi... mit 
Beitrágen der Herren Anton, Dr von dem Busch, 
Dr Dunker, Dr Jonas, Bergrath Koch, Dr Pfeiffer, 
Dr Troschel. Cassel, Theodor Fischer, 1: 1-24. 

FERUSSAC, А. E. J. P. J. F d'A. € С.Р. DE- 
SHAYES, 1819-1851, Histoire Naturelle 
générale et particulière des mollusques ter- 
restres et fluvialiles. Paris, J. B. Baillière, 1 
(Text): i-vii + 1-402; 2 (Text): i-xvi + 1° part: 1- 
184; 2° part: 1-260; 1 (Atlas): 1-24, 2 (Atlas), 
pls. 70-166 + 5 (fossiles). 

FERUSSAC, А. Е. J. P. J. F. ФА., 1827, Cata- 
logue des espèces de mollusques terrestres 
et fluviatiles, recueillies par M. Rang, office. 
de la marine roy., dans um voyage aux grandes 
Indes. Bulletin des Sciences Naturelles et de 
Géologie, [Deuxième Section du Bulletin 
Universel des Sciences et de L’Industrie], 
10(200): 298-307, (284): 408-412. 

GERLACH, J., 1995, The taxonomy and affini- 
ties of the genus Priodiscus (Mollusca; Gas- 
tropoda; Streptaxidae). Journal of Conchology, 
35(4): 357-368. 

GRAY, J. E., 1837, On a new genus of land shells. 
The Magazine of Natural History, (n.s.), 1(9): 
484-487. 


TAXONOMIC REVISION OF THE GENUS HYPSELARTEMON 11 


GUDE, G. K., 1902, A synopsis of the genus 
Streptaxis and its allies. Proceedings of the 
Malacological Society of London, 5(3): 201- 
244. 

GUDE, G. K., 1903, Additions to the genus 
Streptaxis. Proceedings of the Malacological 
Society of London, 5(4): 324-327. 

HIDALGO, J. G., 1870, Catalogue des coquilles 
terrestres recueillis par les naturalistes de la 
commission scientifique espagnole sur divers 
points de I!’ Amerique Méridionale. Journal de 
Conchyliologie, 18(1): 27-70. 

HIDALGO, J. G., 1872, Moluscos Del viaje al 
Pacifico verificado de 1862 a 1865 por uma 
comision de naturalistas enviada por el 
gobierno Español. (parte 1°: univalvos 
terrestres). Madrid, Imprenta de Miguel 
Ginesta, 152 pp. 

HUPE, H., 1857, Mollusques, 3: 1-96, in: F. 
CASTELNAU, ed., Animaux nouveaux ou rares 
recueillis pendant l’expedition dans les parties 
centrales de ГАтепдие du Sud, de Rio de 
Janeiro a Lima et de Lima au Para: par ordre 
du Governement Francais pendant les anées 
1843 a 1847, sous la direction du Conte Francis 
de Castelnau, 7°, Partie 3 (Zoologie). Paris, P. 
Bertrand. 

KOBELT, W., 1905-1906, Die Raublungen- 
schnecken (Agnatha). Zweite Abtheilung: 
Streptaxidae und Daudebardiidae. In: H. C. 
KUSTER & S. CLESSIN, eds., Systematisches 
Conchylien-Cabinet von Martini und Chemnitz, 
1(12b) (2): 211 pp., pls. 42-71, pp. 1-96, pls. 
42-59 (1905); pp. 97-211, pls. 60-71 (1906). 

KOBELT, W., 1910, Katalog der lebenden 
schalentragenden Mollusken der Abteilung 
agnatha. Jahrbücher des Nassauischen 
Vereins für Naturkunde, 63: 138-196. 

LANGE-DE-MORRETES, F., 1949, Ensaio de 
catalogo dos Moluscos do Brasil. Arquivos do 
Museu Paranaense, 7: 5-216. 

PFEIFFER, L., 1848-1877, Monographia 
heliceorum viventium. Lipsiae, E. A. Brockhaus, 
1: 1484 (1848a); 2: 1-594 (1848b); 3: 1-711 
(1853); 4: 1-920 (1859); 5: 1-565 (1868); 6: 
1598 (1868); 7: 1-674 (1876); 8: 1-729 (1877). 

PFEIFFER, L., 1849, Description of a new Helix 
and Streptaxis, from the collection of H. 
Cuming, Esq. Proceedings of the Zoological 
Society of London, 1848(187): 108-109. 

PFEIFFER, L., 1850a, Descriptions of twenty- 
four new species of Helicea, from the collec- 
tion of H. Cuming, Esq. Proceedings of the 
Zoological Society of London, 1849(197): 126- 
131, 

PFEIFFER, L., 1850b [1846-1853], Die 
Schnirkelschnecken nebst den zunächst 
verwandten Gattungen. Zweiter Thiel. In: H. С. 
KÜSTER & S. CLESSIN, eds., Systematisches 
Conchylien-Cabinet von Martini und Chemnitz, 
1(12) (2): 1-290, pls. 67-124. Nürnberg, Bauer 
& Raspe. 

PFEIFFER, L., 1859; see PFEIFFER, L., 1848- 
1877. 

PFEIFFER, L., 1867, Descriptions de deux 
nouvelles espèces de Streptaxis. Journal de 
Conchyliologie, 15(1): 43-44. 


PFEIFFER, L. & $. CLESSIN, 1881, Nomenclator 
heliceorum viventium quo continetur [sic] 
nomina familiae generum et specierum hodie 
cognitarum disposita ex affinitate naturali. 
Casselis, pp. 1-617. ] 

PICORAL, M. & J. W. THOME, 1998, Anatomia 
e microanatomia do sistema reprodutor de 
Rectartemon (Rectartemon) depressus 
(Heynemann) (Mollusca, Gastropoda, 
Streptaxidae): complexo falico. Revista 
Brasileira de Zoologia, 15(1): 101-107. 

PILSBRY,; H.- Ay: 1897; New "Brazilian 
Streptaxidae. Proceedings of the Academy of 
Natural Sciences of Philadelphia, 49: 477-479. 

PILSBRY, H. A., 1907 [1907-1908], in: G. w. 
TRYON, JR. 8 H. А. PILSBRY, eds., Manual of Con- 
chology; Second Series: Pulmonata. Academy 
of Natural Sciences of Philadelphia, 19(73): 1- 
64; (74): 65-128; (75): 129-192. 

PILSBRY, H.A., 1930, South American land and 
freshwarter mollusks: notes and descriptions 
— VII. Proceedings of the Academy of Natural 
Sciences of Philadelphia, 82(7): 355-365. 

RANG, S., 1831, Descriptions des coquilles 
terrestres recueillies pendant un voyage a la 
cote occidentale d’Afrique, et au Brésil. 
Annalles des Sciences Naturelles, 24: 1-63. 

REEVE, L. A., 1851-1854, Monograph of the 
genus Helix. Conchologia Iconica, 7: 210 pls., 
1495 species. London, Reeve. 

REZENDE, H. E. В., P. A. Е. RODRIGUES & J. 
L. BARROS-ARAUJO, 1962, Sôbre o 
Streptaxis contusus (Férrussac, 1821) (Gas- 
tropoda, Pulmonata, Streptaxidae). Memörias 
do Instituto Oswaldo Cruz, 60(3): 337-345. 

RICHARDSON, C. L., 1988, Streptaxacea: cata- 
log of species. Part |. Streptaxidae. Tryonia, 
4(16i): 1-174. 

SALGADO, N. C. & A. C. S. COELHO, 2003, 
Moluscos terrestres do Brasil (gastropodes 
operculados ou nao, exclusive familias 
Veronicellidae, Milacidae e Limacidae). Pp. 
149-189, in: Z. BARRIENTOS & J. MONGE-NAJERA, 
eds., Malacologia Latinoamericana. /nterna- 
tional Journal of Tropical Biology and Conser- 
vation, 51 (Suppl. 3): 536 pp. 

SOLEM, A., 1956, Non-marine Mollusca from 
Salobra, Matto Grosso, Brazil and a collection 
of south Brazilian Artemon. Notulae Nature of 
the Academy of Natural Sciences of Philadel- 
phia, 287: 1-14. 

SOLEM, A., 1974, Patterns of radular tooth struc- 
ture in carnivorous land snails. The Veliger, 
17(2): 81-88. Е 

THIELE, J., 1927, Uber einige brasilianische 
Landsnecken. In: Ergebnisse einer Zoologi- 
schen Forschungsreise in Brasilien 1913-1914 
von Prof. Dr. Е. Bresslau. Abhandlungen der 
Senckenbergischen Naturforschenden 
Gesellschaft, 40(3): 307-329. 

THIELE, J., 1931, Handbuch der systematischen 
Weichtierkunde. Jena, Gustav Fischer, 1(2): 
377-778. 

TRYON, G. W., JR., 1885, Manual of conchol- 
ogy. Second Series: Pulmonata. Conchological 
Section of the Academy of Natural Science of 
Philadelphia, 1: 5-364. 


12 BARBOSA ET AL. 


TRYON, G. W., JR., 1888, Manual of Conchol- 
ogy. Second Series: Pulmonata. Conchological 
Section of the Academy of Natural Science of 
Philadelphia, 4: 3-296. 

VAUGHT, K. C., 1989, A classification of the liv- 
ing Mollusca. Melbourne, Florida, American 
Malacologists.189 pp. 

WENZ, W. v., 1947, Zur Taxonomie der Euthy- 
neura. Archiv für Molluskenkunde, 76(1/3): 36. 

ZILCH, A., 1960, Euthyneura. In: W. WENZ, Gas- 
tropoda. In: O. H. SCHINDEWOLF, Handbuch der 
Paläozoologie. Berlin, Gebrüder Borntraeger, 
2(3): 401-600; (4): 601-834 + xii. 

ZILCH, A., 1961, Die Typen und Typoide des 
Natur-Museums Senckenberg, 24: Mollusca, 
Streptaxidae. Archiv fur Molluskenkunde, 90(1/ 
3): 79-116, pls. 5-9. 


Revised ms. accepted 15 March 2007 


MALACOLOGIA, 2008, 50(1-2): 13-56 


COMPARATIVE HISTOLOGY OF RADULA-SUPPORTING 
STRUCTURES IN GASTROPODA 


Shiho Katsuno?” € Takenori Sasaki** 


ABSTRACT 


A comparative histological study was undertaken to reveal the morphological diversity and 
systematic characters of a radula-supporting organ of gastropods. Observations on 33 
species, all from different families, revealed six major morphological characters: (1) the number 
of odontophoral cartilages or radular bolsters: 0, 1 (fused), 2, 4, 5, 6 and 10; (2) histology 
categorized into 6 types based on the properties of cartilage matrix and cells; (3) the presence 
or absence of an enclosing membrane of the cartilages or radular bolsters; (4) the presence 
or absence of overlapping of the right and left cartilages or radular bolsters; (5) the closest 
position of the cartilages or radular bolsters to each other in cross section at ventral or dorsal 
side; and (6) the insertion areas of the ventral approximator muscle connecting the cartilages 
or radular bolsters — ventral, medial, or outer lateral area. Outgroup and ingroup comparisons 
based on recent phylogenetic hypotheses suggest the following evolutionary scenario for 
gastropod radula-supporting organs: the ancestral gastropod is assumed to have possessed 
two pairs of odontophoral cartilages with a thick matrix and ventrally connected by the 
approximator muscle. The cartilages have possibly independently increased in number in 
patellogastropods and Neritimorpha, decreased into a one pair, single piece or lost in 
Caenogastropoda, and replaced by connective tissue and muscle fibers in Heterobranchia. 
Some taxa such as Cypraeidae have gained a unique histology. The cartilages or radular 
bolsters are closest ventrally in cross section in the majority of gastropods but closest dorsally 
in part of the taenioglossate Caenogastropoda. The diversification of these character states 
in gastropods seems to be phylogenetically constrained, not ecologically. 

Key words: odontophore, odontophoral cartilage, radular bolster, ventral approximator 


muscle, histology, morphological diversity, Gastropoda. 


INTRODUCTION 


Gastropoda has gained the most diversified 
anatomy and ecology among the nine classes 
of the phylum Mollusca and are therefore an 
interesting subject for comparative anatomy 
(e.g., Haszprunar, 1988a; Ponder & Lindberg, 
1997). In phylogenetic studies, morphology- 
based cladistic analyses have been carried out 
using more than 100 anatomical characters for 
gastropods (Salvini-Plawen & Steiner, 1996; 
Ponder & Lindberg; 1997; Sasaki, 1998; Barker, 
2001; Dayrat & Tillier, 2002; Strong, 2003). 
Among them, feeding structures (radular, 
odontophore and related structure) provide a 
number of characters useful for higher phylog- 
eny and are of great importance in compara- 
tive morphology. 


Concerning radula-supporting structure, the 
presence or absence of the cartilages and the 
number of cartilage pairs have been verified 
to define higher taxonomic groups of gastro- 
pods (Salvini-Plawen & Steiner, 1996: char- 
acter #22 for Strepeneura; Ponder & Lindberg; 
1997: #68: Sasaki, 1998: #54-56; Barker, 
2001: #2; Dayrat & Tillier, 2002: #34; Strong, 
2003: #13). Most notably, cartilage-less 
Heterobranchia is contracted by cartilage- 
bearing “prosobranchs” (see above for referen- 
ces). The revision of published data, however, 
suggests greater diversity potentially exists in 
cartilage morphology and histology, but they 
have not been fully investigated across major 
gastropod groups comparatively (see Discus- 
sion for source of previous descriptions). In 
addition, there has been some confusion in the 


‘Department of Earth and Planetary Science, The University of Tokyo, 7-3-1 Hongo Bunkyo-ku, Tokyo, 113-0033, Japan 
“Present address: Eisai Co., Ltd., Koishikawa 4-6-10, Bunkyo-ku, Tokyo 112-8088, Japan 
“The University Museum, The University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033, Japan 


“Corresponding author: sasaki@um.u-tokyo.ac.jp 


14 KATSUNO & SASAKI 


use of terms (odontophore, cartilages, bolsters, 
etc.) and interpretations of homology among 
different molluscan classes. 

In a search for useful systematic characters 
in gastropods, we attempted a comparative 
study of radula-supporting organs. Based on 
new results, we categorize discrete character 
states of various parts, summarize their taxo- 
nomic distribution in gastropods, and discuss 
their systematic implications, homology, and 
relevance to ecology. 


MATERIAL AND METHODS 


Living specimens of 33 species belonging to 
33 families of Gastropoda were collected from 
various environments covering terrestrial, 
fresh- and blackish-water, intertidal to bathyal 
environments in Japan (Table 1). After removal 
from the shell, the soft parts of animals were 
fixed in 4% formalin, preserved in 70% etha- 
nol, and dissected under a binocular micro- 
scope. For histological sectioning, a head 
containing the buccal mass was dehydrated 
with a series of ethanol, embedded in paraffin, 
serially cross-sectioned at the thickness of 4, 
6 or 8 um, and stained by Heidenhain’s Azan 
method in which collagenous tissues are 
stained in blue and muscle fibers are in red. 
Histology of radular-supporting organs was 
observed under a light microscope and photo- 
graphed. Systematics at family or higher level 
in this paper follows Bouchet & Rocroi (2005). 


Abbreviations 


ac anterior cartilage 

alc anterolateral cartilage 
cc cartilage cell 

ct connective tissue 

e  extramatrix substance 
es esophagus 

fc fibrous connective tissue 
ic inner cartilage 

m cartilage matrix 

mc median cartilage 

mf muscle fiber 

n nucleus 

oc odontophoral cartilage 
ouc outer cartilage 

pc posterior cartilage 

pdc posterodorsal cartilage 
ra radula 

rb radular bolster 

rv  radular vesicle 

te tendon-like structure 


va ventral approximator muscle 

vad ventral approximator muscle dorsal layer 
vav ventral approximator muscle ventral layer 
vc ventral cartilage 


Terminology 


Revision and redefinition of the descriptive 
terms for molluscan radular-supporting organs 
are necessary because different terminology 
has been used for similar structures by vari- 
ous authors in the past. We redefine these 
terms below. 

The cartilage or odontophoral cartilage are 
widely used terms (e.g., Fretter & Graham, 
1962: 177; Hubendick, 1978: 13; Salvini- 
Plawen, 1988: 331; Ponder & Lindberg, 1997: 
145; Sasaki, 1998; Sasaki et al., 2006a, b), 
but has been used without definition in most 
cases. In this study, we define the odontophoral 
cartilages in gastropods as follows: rigid 
structure(s) in the odontophore composed 
exclusively of similar-sized turgescent cells 
partitioned by an extracellular matrix. This 
definition is consistent with a general and 
broad definition of invertebrate cartilages 
based on the presence of an extracellular 
matrix (Cole & Hall, 2004). 

The bolster or radular bolster is another 
widely used term (Hyman, 1967: 241; Salvini- 
Plawen, 1988: 329: Scheltema et al., 1994: 
40; Voltzow, 1994: 167; Shimek & Steiner, 
1997). It has not been defined histologically 
but used for any massive structure supporting 
the radula within the odontophore. However, 
to emphasize histological differences from the 
odontophoral cartilages defined above, we use 
this term for any radula-supporting structures 
that possess the identical topology with 
odontophoral cartilages, but lacking true car- 
tilaginous tissue, as in Heterobranchia. 

The odontophore is sometimes used for a 
supporting structure lacking the cartilage (e.g., 
Mackenstedt & Markel, 2001) or synonymous 
with the radular bolster (Voltzow, 1994: 167). 
However, the term is also equivalent to the 
term buccal mass in some literature (e.g., 
Fretter & Graham, 1962: 153). To avoid con- 
fusion, we apply the term odontophore to a 
larger unit containing the radula, radula-sup- 
porting structures (odontophoral cartilages 
and/or radular bolsters) and their connecting 
muscles, as has been widely used (e.g., 
Hyman, 1967; Hubendick, 1978: 13; Salvini- 
Plawen, 1988: 329; Sasaki, 1998). 

The radular vesicle has been applied exclu- 
sively to hollow fluid-filled radula-supporting 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 15 


structures found in Polyplacophora (Wing- 
strand, 1985) and Monoplacophora (Lemche 
& Wingstrand, 1959; Wingstrand, 1985; Schae- 
fer & Haszprunar, 1996; Haszprunar & Schae- 


fer, 1996, 1997). Based on clear differences in 
its inner structure from other similar odonto- 
phoral cartilages, the use of this term should 
be restricted to those two classes. 


TABLE 1. Sampling data of material examined in this study. All localities are in Japan. 


Higher Taxa 


Polyoplacophora 
Ischnochitonidae 


Species 


Ischnochiton comptus 


Locality and Sampling Date 


Misaki, Miura Peninsula, June 3-5, 2004 


Scaphopoda 
Dentaliidae Dentalium octanglatum Koajiro Bay, Miura Peninsula, Nov. 12, 2001 
Gadilinidae Episiphon subrectum Off Katsuura, Wakayama Prefecture, 51-59 m, 


Patellogastropoda 


R/V Tansei-maru, cruise KT-99-17, station 
KU4, Nov. 29, 1999 


Nacellidae Cellana grata Takeoka, Futtsu, Chiba Prefecture, May 25, 2002 
Vetigastropoda 

Haliotidae Haliotis diversicolor aquatilis Fish market in Tokyo, Aug. 1, 2003 

Trochidae Monodonta labio forma con- Jogashima Island, Misaki, Kanagawa Pre- 


Cocculiniformia 
Cocculinidae 


Neritimorpha 
Neritidae 


fusa 


Cocculina sp. cf. japonica 


Nerita albicilla 


Taenioglossate Caenogastropoda 


fecture, June 5, 2004 


Off Hino-Misaki, Shimane Prefecture, 397- 
404 m, R/V Tansei-maru, cruise KT-98-17, 
station 8, Sep. 27-28, 1998 


Jogashima Island, Misaki, Kanagawa Prefec- 
ture, June 5, 2004 


Cyclophoridae Cyclophorus herklotsi Minakawa, Kuroshio, Kouchi Prefecture, April 
24, 1999 

Viviparidae Cipangopaludina chinensis  Ikura, Wakasa, Fukui Prefecture, June 20, 

laeta 2002 

Batillariidae Batillaria cumingii Aburatsubo, Misaki, Kanagawa Prefecture, 
June 5, 2004 

Calyptraeidae Crepidula onyx Kisarazu, Chiba Prefecture, June 2003 

Cypraeidae Cypraea boivinii Aburatsubo, Misaki, Kanagawa Prefecture, 
Aug. 3, 2003; Banda, Tateyama, Chiba Pre- 
fecture, Sep. 27-28, 1998 

Littorinidae Littorina brevicula Jogashima Island, Misaki, Kanagawa Prefec- 
ture, April 24, 2004; Odaiba, Tokyo, April 28, 
2004 

Naticidae Glossaulax didyma Kisarazu, Chiba Prefecture, June, 2003 

Assimineidae Assiminea japonica Mihama, Fukui Prefecture, June 20, 2002 

Strombidae Strombus luhuanus Uehara, Iriomote Island, Okinawa, May 10, 
2004 

Ranellidae Fusitriton oregonensis Otsuchi, lwate Prefecture, July 15, 1999 

Hipponicidae Hipponix conica Fish market in Tokyo, Aug. 20, 2003 

Vermetidae Serpulorbis imbricatus Jogashima Island, Misaki, Kanagawa Pre- 


fecture, May 20, 2003 


(continues) 


16 KATSUNO & SASAKI 


(continued) 
Higher Taxa Species Locality and Sampling Date 
Neogastropoda 
Buccinidae Japeuthria ferrea Jogashima Island, Misaki, Kanagawa Pre- 


fecture, April 24, 2004 


Columbellidae 


Nassariidae 
Muricidae 


Mitridae 


Conidae 


Heterobranchia 
Haminoeidae 


Philinidae 


Cylichnidae 


Cavoliniidae 
Aplysiidae 


Chromodorididae 


Siphonariidae 


Lymnaeidae 
Onchidiidae 


Bradybaenidae 


Euplica scripta 


Nassarius festiva 
Thais clavigera 


Strigatella zebra 


Conus ebraeus 


Haloa japonica 
Philine argentata 


Nipponoscaphander japonica 


Cavolinia uncinata 
Aplysia oculifera 


Hypselodoris festiva 


Siphonaria japonica 


Lymnaea stagnalis 
Peronia sp. cf. verruculatum 


Acusta despecta sieboldiana 


Jogashima Island, Misaki, Kanagawa Pre- 
fecture, June, 2003 

Kisarazu, Chiba Prefecture, June, 2003 

Jogashima Island, Misaki, Kanagawa Pre- 
fecture, April 24, 2004 

Hoshizunano-hama, Iriomote Island, Okinawa, 
May 13, 2004 

Hoshizunano-hama, Iriomote Island, Okinawa, 
May 13, 2004 


Jogashima Island, Misaki, Kanagawa Pre- 
fecture, May 20, 2003 

Lake Hamana, Shizuoka Prefecture, Aug. 
1996 

Off Katsuura, Wakayama Prefecture, 99-102 
т, R/V Tansei-maru, cruise KT-99-17, sta- 
tion KU5, Nov. 30, 1999 

Off Izu Islands, R/V Shinyo-maru sta. 1998-L1, 
Oct. 19, 1998 

Jogashima Island, Misaki, Kanagawa Pre- 
fecture, May 20, 2003 

Arasaki, Yokosuka, Kanagawa Prefecture, 
June 23, 2002; Jogashima Island, Misaki, 
Kanagawa Prefecture; Aug. 14, 2003; Abu- 
ratsubo, Misaki, Kanagawa Prefecture; 
June 3-5, 2004 

Jogashima Island, Misaki, Kanagawa Pre- 
fecture, April 24, 2004; Jogashima Island, 
Misaki, Kanagawa Prefecture, Aug. 1, 2003 

Aqualium in The University of Tokyo, 2003 

Aburatsubo, Misaki, Kanagawa Prefecture, 
Aug. 1, 2003 

Omachi, Nagano Prefecture, July 2002 


RESULTS 


Clade Patellogastropoda 
Superfamily Nacelloidea 
Family Nacellidae 


Cellana grata (Gould, 1859) 
(Figs. 1A, 2) 


Odontophore containing five pairs of 
cartilages — anterior (ac), anterolateral (alc), 
ventral (vc), posterodorsal (pdc), posterior 


cartilages (pc) (Figs. 1A, 2A-C). Anterior and 
anterolateral cartilages larger than posterior 
ones (Fig. 1A), former especially elongate. 
Anterolateral cartilages attached to anterolat- 
eral side of anterior cartilages, ventral 
cartilages to anteroventral side. Posterior 
cartilages overlying posterior end of anterior 
cartilages; posterodorsal cartilages closely 
anterior to posterior cartilages. All cartilages 
except anterior one show similar histology 
having stiffened matrix (m) and cartilage cells 
(cc) containing small nuclei (n) that tend to be 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 17 


J Sele, ee 


FIG. 1. Outline of odontophoral cartilages. A: Cellana grata (Nacellidae); B: Haliotis diversicolor 
aquatilis (Haliotidae); C: Nerita albicilla (Neritidae); D: Cipangopaludina chinensis laeta 
(Cyclophoridae); Е: Littorina brevicula (Littorinidae); Е: Crepidula onyx (Calyptraeidae); С: Cypraea 
boivinii (Cypraeidae); H: Thais clavigera (Muricidae); |: Strigatella zebra (Mitridae); J: Peronia sp. 
cf. verruculatum (Onchidiidae); K: Acusta despecta sieboldiana (Bradybaenidae). All dorsal views. 


18 KATSUNO & SASAKI 


FIG. 2. Cross sections of buccal mass and odontophoral cartilages of Cellana grata (Nacellidae). A: 
Anterior part of buccal mass; B: Middle part of buccal mass; C: Posterior part of buccal mass; D: 
Enlarged view of anterior cartilage; E: Dorsal and ventral layers of ventral approximator muscle; F: 
Enlarged view of anterolateral cartilage. 


located close to matrix (Fig. 2F). Cells of an- nected by two-layered ventral approximator 
terior cartilages smaller and denser than those muscle (Fig. 2A, E); dorsal layer (vad) insert- 
of other cartilages (Fig. 2D). Anterior cartilages ing at anteromedian area of anterior cartilages 
not fused or overlapped, closest to each other (Fig. 2A); ventral layer (vav) connecting ven- 
ventrally in cross section (Fig. 2A-C), con- tral plane of anterior cartilages (Fig. 2A, B). 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 19 


FIG. 3. Cross sections of buccal mass and odontophoral cartilages of Haliotis diversicolor aquatilis 
(Haliotidae). A: Middle part of buccal mass; B: Posterior part of buccal mass; C: Enlarged view of 
ventral approximator muscle; D: Enlarged view of anterior cartilage. 


Clade Vetigastropoda 
Superfamily Haliotoidea 
Family Haliotidae 


Haliotis diversicolor aquatilis Reeve, 1846 
(Figs. 1B, 3) 


Odontophore containing two pairs of car- 
tilages — anterior and posterior cartilages (ac, 
pc: Fig. 3A, B). Posterior cartilages smaller than 
anterior cartilages, adhering to posterior end 
of anterior cartilages (Fig. 1B: see Sasaki, 
1998: fig. 31, for gross morphology). Both 
cartilages consisting of robust matrix (m), car- 
tilage cells (cc) containing small nuclei (n) fre- 
quently attached tightly to matrix (Fig. 3D). 
Anterior cartilages not fused or overlapped, 
facing each other most closely ventrally in 
cross section, connected with tendon-like 
structure (te) (Fig. 3A) and thin layer of ven- 
tral approximator muscle (va) (Fig. 3A, C). 


Superfamily Trochoidea 
Family Trochidae 


Monodonta labio confusa Tapparone- 
Canefri, 1874 


(Fig. 4) 


Odontophore containing two pairs of car- 
tilages, viz. anterior and posterior cartilages (ac, 
pc: Fig. 4A, B). Both cartilages composed of 
stout matrix (m), cartilage cells (cc), with small 
nuclei (n) located close to matrix (Fig. 4D). 
Posterodorsal parts of anterior cartilages con- 
taining characteristic dense zone of thin matrix 
and cells (asterisk in Fig. 4A; magnified in Fig. 
4C). Anterior cartilages not fused or overlapped, 
closest ventrally in cross section, connected by 
ventral approximator muscle (va) including fi- 
brous connective tissue (Fig. 4A). Ventral 
approximator muscle inserting at ventral side 
of anterior cartilages (Fig. AA). 


20 KATSUNO & SASAKI 


FIG. 4. Cross sections of buccal mass and odontophoral cartilages of Monodonta labio confusa 
(Trochidae). A: Anterior part of buccal mass. Asterisk indicates a mass of smaller cells; B: Posterior 
part of buccal mass; C: Enlarged view of part of anterior cartilage marked by asterisk in Fig. A; D: 


Enlarged view of anterior cartilage. 


Clade Cocculiniformia 
Superfamily Cocculinoidea 
Family Cocculinidae 


Cocculina sp. cf. japonica Dall, 1907 
(Fig. 5A, B) 


Odontophore containing single pair of car- 
tilages (oc: Fig. 5A) consisting of cartilage cells 
(cc) and matrix (m). Cartilage matrix consist- 
ing of two elements: (1) thick sheets of matrix 
at nearly constant spacing that divide 
cartilages into multiple horizontal sectors of 
nearly equal size; (2) extremely thin matrix 
connecting horizontal sheets obliquely or ver- 
tically (Fig. 5B). Nuclei (n) of cartilage cells 
relatively large, mostly attached to matrix (Fig. 
SB). Cartilages not fused along entire length, 
closest to each other ventrally without over- 
lapping, and connected by connective tissue 
(ct) and by ventral approximator muscle (va) 
inserting at ventral side of cartilages (Fig. 5B). 


Clade Neritimorpha 
Superfamily Neritoidea 
Family Neritidae 


Nerita albicilla Linnaeus, 1758 
(Figs. 1C, 5C-F) 


Odontophore containing two pairs of car- 
tilages (Fig. 1C) and single unpaired cartilage 
(Fig. 5C, F). Anterior and posterior cartilages 
composed of thick cartilage matrix (m), carti- 
lage cells (cc), very small nuclei (n) beside 
matrix (Fig. 5D). Median cartilage showing 
similar histology but its matrix thinner than that 
of other cartilages (Fig. 5D). Anterior cartilages 
(ac) longitudinally elongate; posterior car- 
tilages (pc) much smaller, attached to poste- 
rior end of anterior cartilages; median cartilage 
(mc) thin, club-shaped, inserted between an- 
terior cartilages (Fig. 1C). Anterior cartilages 
unfused, closest ventrally without overlapping 
in cross section, connected at ventral level by 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 21 


FIG. 5. Cross sections of buccal mass and odontophoral cartilages. A, B. Cocculina sp. cf. japonica 
(Cocculinidae). A: Section in middle part of buccal mass; B: Enlarged view of Fig. 5A to show histol- 
ogy of cartilages, ventral approximator muscle (va) and connective tissue (ct); C, F. Nerita albicilla 
(Neritidae). C: Section in middle part of buccal mass; D: Enlarged view of anterior cartilage; E: En- 
larged view of ventral approximator muscle; F: Median cartilage. 


ventral approximator muscle (va) inserting at Clade Caenogastropoda 

ventral side (Fig. 5C). Informal group Architaenioglossa 
Sasaki (1998: p. 111, fig. 75) described the Superfamily Cyclophoroidea 

median cartilage as a pair of symmetrical el- Family Cyclophoridae 

ements. However, the median cartilage is ac- 

tually a single body in histological observa- Cyclophorus herklotsi Martens, 1861 


tions. (Fig. 6A, B) 


22 KATSUNO & SASAKI 


FIG. 6. Cross sections of buccal mass and odontophoral cartilages. A, B. Cyclophorus herklotsi 
(Cyclophoridae). A: Section in anterior part of buccal mass; B: Enlarged view of odontophoral carti- 
lage; C, D. Cipangopaludina chinensis laeta (Viviparidae). C: Section in middle part of buccal mass; 


D: Enlarged view of odontophoral cartilage. 


Odontophore containing single pair of un- 
fused cartilages (oc) (Figs. 1D, 6A) with thin 
matrix (m), cartilage cells (cc), and nuclei (n) 
that are large in size relative to cartilage cells, 
located in various parts within them (Fig. 6B). 
Cartilages closest ventrally, connected by ven- 
tral approximator muscle (va) inserting at outer 
lateral side (Fig. 6A). Left cartilage partially 
overlying right cartilages. 


Superfamily Viviparoidea 
Family Viviparidae 


Cipangopaludina chinensis laeta (Martens, 
1860) 
(Figs. 1D, 6C, D) 


Odontophore containing single pair of 
unfused cartilages (oc: Fig. 6C) with thin ma- 
trix (m), cartilage cells (cc), relatively large 


nuclei (n) mostly in contact with matrix (Fig. 
6B). Cartilages closest and connected ven- 
trally by single-layered ventral approximator 
muscle (va) inserting at outer lateral side (Fig. 
6C). Left cartilage underlying right cartilage. 


Clade Sorbeoconcha 
Superfamily Cerithioidea 
Family Batillariidae 


Batillaria cumingii (Crosse, 1862) 
(Fig. 7A, B) 


Odontophore containing single pair of unfused 
cartilages (oc) consisting of thin matrix (m), car- 
tilage cells (cc) with relatively large nuclei (n) 
scattered in various areas inside of cells (Fig. 
7B). Pair of cartilages overlapping ventrally, con- 
nected by ventral approximator muscle (va) in- 
serting at outer lateral side of cartilages (Fig. 7A). 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 23 


FIG. 7. Cross sections of buccal mass and odontophoral cartilages. A, B. Batillaria cumingii 
(Batillariidae). A: Outer lateral origin of ventral approximator muscle; B: Enlarged view of central part 
of odontophoral cartilage; C, D. Crepidula onyx (Calyptraeidae). C: Anterior part of buccal mass; D: 


Enlarged view of odontophoral cartilage. 


Clade Hypsogastropoda 
Clade Littorinimorpha 
Superfamily Calyptoraeoidea 
Family Calyptraeidae 


Crepidula onyx С. В. Sowerby I, 1824 
(Figs, 1F, 70D) 


Odontophore containing single unfused pair 
of cartilages (oc: Figs. 1F, 7C) with thin matrix 
(m), cartilage cells (cc) and relatively large 
nuclei (n) located in various areas within car- 
tilage cells (Fig. 7D). Cartilages closest ven- 
trally without overlapping, connected by ventral 
approximator muscle (va) inserting at outer 
side of cartilages (Fig. 7C). 


Superfamily Cypraeoidea 
Family Cypraeidae 


Cypraea boivinii Kiener, 1843 
(Figs. 1G, 8) 


Odontophore containing single unfused pair 
of cartilages (oc: Figs. 1G, 8A, B) consisting 
of markedly thick bubble-shaped matrix (m), 
with extramatrix substance (e) filling spaces 
between cartilage matrices, cartilage cells (cc) 
with small nuclei (n) located at various areas 
inside of cells (Fig. 8D). Matrix and extramatrix 
substance stained in blue and in red, respec- 
tively. Cartilages closest ventrally without 
overlapping and connected by ventral 
approximator muscle (va) inserting at ventral 
side of cartilages (Fig. 8A, C). 


Superfamily Littorinoidea 
Family Littorinidae 


Littorina brevicula (Philippi, 1844) 
(Figs. 1E, 9A, B) 


Odontophore containing single unfused pair 
of cartilages (oc: Figs. 1E, 9A), consisting of 
thin matrix (m), cartilage cells (cc), large nuclei 


24 KATSUNO & SASAKI 


FIG. 8. Cross sections of buccal mass and odontophoral cartilages of Cypraea boivinii (Cypraeidae). 
A: Anterior part of odontophoral cartilage; B: Posterior part of odontophoral cartilage; C: Ventral 
approximator muscle; D: Enlarged view of odontophoral cartilage. 


(n) in variable positions within cartilage cells (Fig. 
9B). Cartilages closest ventrally, connected by 
ventral approximator muscle (va) inserting at 
outer lateral sides of cartilages (Fig. 9A, B). Left 
cartilage overlying right cartilage (Fig. 9A). 


Superfamily Naticoidea 
Family Naticidae 


Glossaulax didyma (Réding, 1798) 
(Fig. 9C, D) 


Odontophore containing single unfused pair 
of cartilages (oc) consisting of thin matrix (m), 
cartilage cells (cc), and large nuclei (n) occur- 
ring in variable positions within cells (Fig. 9D). 
Cytoplasm of some cartilage cells stained in 
mazenda (Fig. 9D). Cartilages lying closest 
dorsally; right cartilage overlying left one (Fig. 
9C). Ventral approximator muscle not clearly 
identified. 


Superfamily Rissooidea 
Family Assimineidae 


Assiminea japonica Martens, 1877 
(Fig. 10A, B) 


Odontophore containing single unfused pair 
of cartilages (oc: Fig. 10A), consisting of thin 
matrix (m), cartilage cells (cc) with large nu- 
clei (п) attached tightly to matrix (Fig. 10B). 
Matrix and cartilage cells stained in blue and 
faintly in mazenda, respectively. Paired 
cartilages closest ventrally, without overlap- 
ping. Connection between cartilages not ob- 
served. 


Superfamily Stromboidea 
Family Strombidae 


Strombus luhuanus Linnaeus, 1758 
(Fig: 106.0) 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 25 


FIG. 9. Cross sections of buccal mass and odontophoral cartilages. A, В. Littorina brevicula 
(Littorinidae). A: Middle part of buccal mass; B: Enlarged view of odontophoral cartilage; C, D. 
Glossaulax didyma (Naticidae). C: Middle part of odontophoral cartilage; D: Enlarged view of 


odontophoral cartilage. 


Odontophore containing single pair of 
unfused cartilages (oc), composed of thin 
matrix (m), cartilage cells (cc), and large nu- 
clei (n) distributed in variable positions in car- 
tilage cells (Fig. 10D). Cartilages closest 
dorsally in cross section. Ventral approximator 
muscle not found. Left cartilage overlying right 
one. 


Superfamily Tonnoidea 
Family Ranellidae 


Fusitriton oregonensis (Redfield, 1846) 
(Fig. 11) 


Odontophore containing single unfused pair 
of cartilages (oc: Fig. 11A, B), consisting of 
thin matrix (m), cartilage cells (cc) with large 
nuclei (n) in variable positions in cells (Fig. 
11D). Pair of cartilages closest ventrally, con- 
nected by ventral approximator muscle (va) 
inserting at inner medial side of cartilages (Fig. 


11C). Left cartilage overlying right cartilage 
(Fig. 11A, B). 


Superfamily Vanikoroidea 
Family Hipponicidae 


Hipponix conica (Schumacher, 1817) 
(Fig. 12A, B) 


Odontophore containing single unfused pair 
of cartilages (oc: Fig. 12A), consisting of thin 
matrix (m), cartilage cells (cc) with large nu- 
clei (n) located on interior wall of matrix (Fig. 
12B). Cartilages closest on dorsal side; left 
cartilage underlying right one (Fig. 12A). Ven- 
tral approximator muscle not found. 


Superfamily Vermetoidea 
Family Vermetidae 


Serpulorbis imbricatus (Dunker, 1860) 
(rig tac. i) 


26 


KATSUNO & SASAKI 


FIG. 10. Cross sections of odontophoral cartilages. A, B. Assiminea japonica 
(Assimineidae). A: Posterior part of odontophoral cartilage; B: Enlarged view of 
odontophoral cartilage; C, D: Strombus luhuanus (Strombidae); C: Odontophoral car- 
tilage and surrounding muscle; D: Enlarged view of odontophoral cartilage. 


FIG. 11. Cross sections of buccal mass and odontophoral cartilage of Fusitriton 
oregonensis (Ranellidae). A: Anterior middle part of buccal mass; B: Posterior middle 
part of buccal mass; C: Tissue of ventral approximator muscle; D: Enlarged view of 


odontophoral cartilage. 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 27 


FIG. 12. Cross sections of buccal mass and odontophoral cartilages. A, B. Hipponix 
conica (Hipponicidae; A, B). A: Middle part of buccal mass; B: Enlarged view of 
odontophoral cartilage; C, D. Serpulorbis imbricatus (Vermetidae). C: Middle part of 
buccal mass; D: Enlarged view of odontophoral cartilage. 


ii 50 um 


FIG. 13. Cross sections of buccal mass and odontophoral cartilages of Japeuthria 
ferrea (Buccinidae). A: Anterior part of buccal mass; B: Posterior part of buccal mass; 
C: Anterior end of odontophoral cartilage; D: Anterior part of odontophoral cartilages 
with ventral approximator muscle. 


28 KATSUNO & SASAKI 


Odontophore containing single pair of 
unfused cartilages (oc: Fig. 12C), thin matrix 
(m), cartilage cells (cc), large nuclei (n) in vari- 
able positions within cartilage cells (Fig. 12D). 
Cartilages closest ventrally, connected by ven- 
tral approximator (va) muscle inserting at outer 
lateral side of cartilages (Fig. 12C). Left carti- 
lage overlying right cartilage anteriorly. 


Clade Neogastropoda 
Superfamily Buccinoidea 
Family Buccinidae 


Japeuthria ferrea (Reeve, 1847) 
(Fig, 13) 


Odontophore containing one fused pair of 
cartilage (oc: Fig. 13A, B), consisting of thick 
matrix (m), cartilage cells (cc), large nuclei (n) 
that tend to be located in middle of cells (Fig. 
13C, D). Cartilage U-shaped, symmetrical, 


connected anteriorly between right and left 
(Fig. 13A, C). In posterior area, branches of 
cartilage separate, connected ventrally by ven- 
tral approximator muscle (va) inserting at ven- 
tral side of cartilage (Fig. 13D). Right and left 
elements not overlapping each other. 


Family Columbellidae 


Euplica scripta (Lamarck, 1822) 
(Fig. 14) 


Odontophore containing single pair of 
cartilages (oc) fused anteriorly but separated 
into right and left branches posteriorly without 
overlapping (Fig. 14A, B), composed of thick 
matrix (m), cartilage cells (cc), and large nu- 
clei (п) situated centrally within cells (Fig. 14С). 
Right and left branches of cartilage connected 
by ventral approximator muscle (va) inserting 
at ventral area of cartilages (Fig. 14B, D). 


FIG. 14. Cross sections of buccal mass and odontophoral cartilages of Euplica scripta (Columbellidae). 
A: Anterior part of buccal mass; B: Middle part of buccal mass; C: Enlarged view of odontophoral 
cartilage; D: Middle part of anterior cartilage and ventral approximator muscle. 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 29 


Family Nassariidae 


Nassarius festiva (Powys, 1833) 
(Figs 15) 


Odontophore containing single pair of 
cartilages fused anteriorly (oc: Fig. 15A) but 
separated posteriorly without overlapping (Fig. 
15B, C). Histologically cartilage consisting of 
thick matrix (m), cartilage cells (cc), and large 
nuclei (n) located centrally within cells (Fig. 15D). 
Right and left elements of cartilage connected 
by ventral approximator muscle (va) inserting 
at ventral side of cartilages (Fig. 15B, D). 


Superfamily Muricoidea 
Family Muricidae 


Thais clavigera (Kuster, 1860) 
(Figs. 1H, 16A-D) 


Odontophore containing single cartilage 
(oc) which consists of two extremely elon- 
gate branches fused anteroventrally (Figs. 
1H, 16C), strikingly curved outwards with- 
out overlapping (Fig. 1H). Tissue of 
cartilages composed of thick matrix (m), 
cartilage cells (cc), and large nuclei (n) situ- 
ated centrally within cartilage cells (Fig. 16C, 
D). Right and left elements of cartilage con- 
nected by ventral approximator muscle (va) 
inserting at ventral area of cartilage (Fig. 
16A, D). 


Family Mitridae 


Strigatella zebra (Lamarck, 1811) 
(Figs. 11, t6E. F) 


Odontophore containing single unfused 
pair of cartilages (oc: Figs. 11, 16E) that are 


FIG. 15. Cross sections of buccal mass and odontophoral cartilages of Nassarius festiva (Nassariidae). 
A: Anterior end of odontophoral cartilage; B: Posterior middle part of buccal mass; C: Posterior part 
of buccal mass; D: Enlarged view of odontophoral cartilage. 


30 KATSUNO & SASAKI 


composed of thick extracellular matrix (m), 
cartilage cells (cc) and large nuclei (n) tend- 
ing to occur in center of cartilage cells (Fig. 
16F). Pair of cartilages closest ventrally with- 
out overlapping (Fig. 16E), connected by ven- 
tral approximator muscle (va) inserting at 
ventral area (Fig. 16F). 


Superfamily Conoidea 
Family Conidae 


Conus ebraeus Linnaeus, 1758 


Buccal mass present as muscular bulb at base 
of proboscis, but odontophoral cartilage absent. 


FIG. 16. Cross sections of buccal mass and odontophoral cartilages. A-D. Thais clavigera (Muricidae). 
A: Middle part of buccal mass; B: Posterior part of buccal mass; C: Anterior end of odontophoral 
cartilages; D: Enlarged view of odontophoral cartilages and ventral approximator muscle; E, F. 
Strigatella zebra (Mitridae). E: Middle part of buccal mass; F: Enlarged view of odontophoral cartilages 
and ventral approximator muscle. 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA a 


Clade Heterobranchia Haloa japonica (Pilsbry, 1895) 
Informal group Opisthobranchia (Fig. 17A, B) 
Clade Cephalaspidea 
Superfamily Haminoeoidea Odontophore lacking cartilage or membrane- 
Family Haminoeidae enclosed radular bolster (Fig. 17A; see Figs. 


FIG. 17. Cross sections of buccal mass. A, B. Haloa japonica (Haminoeidae). A: Posterior part of 
buccal mass; B: Enlarged view of radular bolster; C, D. Philine argentata (Philinidae). C: Anterior part 


of buccal mass; D: Enlarged view of part of Fig. 17A; E, F. Nipponoscaphander japonica (Cylichnidae). 
E, F: Enlarged view of connective tissue and muscle fibers in radular bolster. 


32 KATSUNO & SASAKI 


19-22 for comparison), densely filled with fi- 
brous connective tissue and muscle fibers ar- 
ranged in variable directions (Fig. 17B). Ventral 
approximator muscle not observed. 


Superfamily Philinoidea 
Family Philinidae 


Philine argentata Gould, 1859 
(Fig. 17C, D) 


Odontophore lacking true cartilage or mem- 
brane-enclosed radular bolster (Fig. 17C), 
filled with loosely organized network-like struc- 
ture (Fig. 17D), being composed of fibrous 


FIG. 18. Cross sections of buccal mass. A, B: Cavolinia uncinata (Cavoliniidae). A: Middle part of 
buccal mass; B: Enlarged view of part of Fig. 18A; C, D. Aplysia oculifera (Aplysiidae). C: Middle part 
of buccal mass; D: Enlarged view of connective tissue and muscle fibers in radular bolster; E, F. 
Hypselodoris festiva (Chromodorididae). E: Middle part of buccal mass; F: Enlarged view of radular 
bolster (right) and surrounding muscles (left) separated by a sheet of collagen fibers (asterisk). 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 33 


connective tissue and muscle fibers. Ventral 
approximator muscle not identified. 


Family Cylichnidae 


Nipponoscaphander japonica (A. Adams, 
1862) 
(Fig. LAB Ar) 


Odontophore lacking cartilage or membrane- 
enclosed radular bolster, filled densely with 
complex of regularly radiating fibrous connec- 
tive tissue and muscle fibers uniformly (Fig. 
17E, F). Ventral approximator muscle not ob- 
served. 


Clade Thecosomata 
Superfamily Cavolinioidea 
Family Cavoliniidae 


Cavolinia uncinata (Rang, 1829) 
(Fig. 18A, B) 


Odontophore lacking cartilage or membrane- 
enclosed radular bolster, containing remarkably 
loose complex of fibrous connective tissue, 
filled internally with large space of blood sinus, 
surrounded exteriorly by layer of epithelial cells 
(Fig. 18A, В). Ventral approximator muscle not 
identified. 


Clade Aplysiomorpha 
Superfamily Aplysioidea 
Family Aplysiidae 


Aplysia oculifera A. Adams & Reeve, 1850 
(Fig. 18C, D) 


Odontophore lacking true cartilaginous tis- 
sue (Fig. 18C), filled with complex of fibrous 
connective tissue (fc) and muscle fibers (mf). 
Middle portion of radular bolsters occupied by 
bubble-shaped spaces of variable size, small 
granular nuclei (n) widely scattered in fibrous 
connective tissue (Fig. 18D). Dorsal and ven- 


FIG. 19. Cross sections of buccal mass and radular bolsters of Siphonaria japonica (Siphonariidae). 
A: Middle part of buccal mass; B: Anterior end of radular bolster; C: Middle part of radular bolsters 
connected by ventral approximator muscle (va); D: Enlarged view of radular bolster. 


34 KATSUNO & SASAKI 


FIG. 20. Cross sections of buccal mass and radular bolsters of Lymnaea stagnalis (Lymnaeidae). A: 
Anterior part of buccal mass; B: Middle part of buccal mass; C: Anterior part of radular bolster; D: 


Enlarged view of radular bolster; E: Posterior part of buccal mass; F: Connection between ventral 
approximator muscle and radular bolster. 


tral portions of radular bolsters lacking bubble- Clade Euctenidiacea 
like tissue and densely packed with connec- Subclade Doridacea 
tive and muscular tissue. Ventral approximator Superfamily Doridoidea 


muscle not observed. Family Chromodorididae 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA Ste) 


Hypselodoris festiva (A. Adams, 1861) 
(Fig. ASEAR) 


Odontophore containing paired stiffened 
radular bolsters (ro: Fig. 18E) enwrapped by 
thin fibrous layer (asterisk in Fig. 18F), sharply 
separated from other buccal musculature, but 
fused with radular retractor muscle ventrally 
(Fig. 18E). Radular bolsters filled with densely 
packed fibers (Fig. 18F). Ventral approximator 
muscle not identified. 


Informal group Pulmonata 
Informal group Basommatophora 
Superfamily Siphonarioidea 
Family Siphonariidae 


Siphonaria japonica (Donovan, 1824) 
(719.19) 


Odontophore containing fused pair of radu- 
lar bolsters (rb: Fig. 19A), not containing car- 
tilage-like structure. Histologically radular 


bolsters consisting of variously oriented mix- 
ture of fibrous connective tissue (fc), muscle 
fibers (mc), darkly stained granular nuclei, 
densely scattered small transparent spaces 
that were not stained by Azan method. 

Right and left elements of radular bolsters 
fused in anterior end (Fig. 19B), closest ven- 
trally without overlapping in cross section (Fig. 
19A), connected by ventral approximator 
muscle (va) inserting at ventral area of radu- 
lar bolsters (Fig. 19C). Radular bolsters sepa- 
rated from surrounding buccal musculature by 
membrane (Fig. 19A). 


Clade Hygrophila 
Superfamily Lymnaeoidea 
Family Lymnaeidae 


Lymnaea stagnalis (Linnaeus, 1758) 
(Fig. 20) 


Odontophore containing fused pair of radu- 
lar bolsters (rb: Fig. 20B) which are composed 


FIG. 21. Cross sections of buccal mass and radular bolster of Peronia sp. cf. verruculatum 
(Onchidiidae). A: Anterior end of buccal mass; B: Middle part of buccal mass; C: Enlarged view of 
radular bolster; D: Connection between ventral approximator muscle and radular bolster. 


36 KATSUNO & SASAKI 


histologically of loose fibrous connective tis- 
sue (fc) and muscle fibers (mf) (Fig. 20D). 
Cartilage matrix absent. Right and left ele- 
ments of radular bolster fused at anterior end 
(Fig. 20A, C), closest ventrally without over- 
lapping, and connected by ventral approx- 
imator muscle (va) inserting at ventral side 
(Fig. 20B). Tissue of radular bolsters not 
merged with outer buccal musculatures by its 
enclosing membrane (Fig. 20A, B). 


Clade Eupulmonata 
Clade Systellommatophora 
Superfamily Onchidioidea 

Family Onchidiidae 


Peronia sp. cf. verruculatum (Cuvier, 1830) 
(Figs. 14.21) 


Odontophore containing unfused pair of 
radular bolsters (rb: Figs. 1J, 21A, B) consist- 
ing of fibrous connective tissue (fc) and muscle 
fibers (mf: Fig. 21C). Cartilage matrix absent. 
Pair of radular bolsters closest ventrally with- 
out overlapping and connected by single layer 


of ventral approximator muscle (va: Fig. 21D). 
Ventral approximator muscle inserting at ven- 
tral area of radular bolsters (Fig. 21A, B). 
Radular bolsters clearly delimited from outer 
buccal musculature by membrane (Fig. 21B). 


Clade Stylommatophora 
Informal group Sigmurethra 
Superfamily Helicoidea 
Family Bradybaenidae 


Acusta despecta sieboldiana (Pfeiffer, 1850) 
(2195. IK, 22) 


Odontophore containing fused paired radu- 
lar bolster (rb: Figs. 1K, 22A), composed of 
condensed fibrous connective tissue and 
muscle fibers (Fig. 22D). Cartilage matrix ab- 
sent. Right and left elements of radular bol- 
ster fused at anterior end, closest ventrally 
without overlapping in cross section (Fig. 22C), 
and connected by ventral approximator muscle 
(va) on ventral side (Fig. 22C). Radular bol- 
ster sharply delimited from surrounding buc- 
cal musculature by membrane (Fig. 22C). 


FIG. 22. Cross sections of buccal mass and radular bolsters of Acusta despecta sieboldiana 
(Bradybaenidae). A: Anterior end of buccal mass; B: Anterior end radular bolster; C: Connection be- 
tween ventral approximator muscle (va) and radular bolster (rb); D: Enlarged view of radular bolster. 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 37 


TABLE 2. Taxonomic distribution of odontophoral cartilages or radular bolsters. ++ = present in all 
species examined; + = present in part; - = absent. Co: Cocculinoidea, He: Heterobranchia, Ng: 
Neogastropoda, Ne: Neritimorpha, Pa: Patellogastropoda, tC: taenioglossate Caenogastropoda, Ve: 
Vetigastropoda. See caption of Appendix for abbreviations of cartilages. 

“Anterior cartilage is type 1, and median cartilage is type 3. 

**Dorsal layer originating from medial side, and ventral layer from ventral side. 


Pa Ve Co Ne tC Ng He 
Number of odontophoral 0 - - - E o ai ir 
cartilages or radular bolsters 1 (fused) - - = : = + + 
2 - + ++ ь ++ + + 
4 (ac + alc) + = E € 5 2 E 
4 (ac + adc) - of - - - - - 
4 (ac + pc) - + - - - - - 
5 г + = ++ - - - 
6 Ñ $ i 3 г Е 
10 - - = ‘ + A 
Composition of ac ++ ++ ++ ++ ++ ++ A 
odontophoral cartilages alc ++ : 2 3 = e : 
pc + = ++ = = - 
adc - - E : > a 
pdc + E ы к = _ L 
mc - + 5 ++ : a 3 
Histology Type 1 ++ + L ++* a s s 
Type 2 - ++ 5 - A x 
Type 3 - 2 eu Е 2 
Type 4 - - 3 . : 2 
Type 5 à á E d d a h 
Type 6 - я a : e J E 
Membrane enclosing carti- ++ ++ ++ ++ ++ ++ + 
lages or radular bolsters 
Overlaping of right and left 4 = A > fe L 
main cartilages or radular 
bolsters 
Closest position of main ventral side ++ ++ ++ ++ + ++ ++ 
cartilages ог radular bolsters dorsal side - - - : + e Е 
Insertion of ventral ventral side +** ++ ++ ++ + ++ = 
approximator muscle medial side qe - - 5 + . x 
outer lateral side - - - - + : . 
dorsal side - - 5 r : o Е 


absent - = à er e E 


38 KATSUNO & SASAKI 


DISCUSSION 
Morphological Diversity 


Greater morphological diversity in gastropod 
radula-supporting organs was revealed in this 
study than previously acknowledged. The char- 
acters mostly used for phylogenetic analysis 
are the number of cartilages and the presence 
or absence of the cartilages (Haszprunar 


1988a: table 2; Ponder & Lindberg, 1997: char- 
acter #68; Sasaki, 1998: characters #54—56; 
Barker, 2001: character #2; Dayrat & Tillier, 
2002: character #34; Strong, 2003: character 
#13). 

In this study, notable differences were ob- 
served in the following six characters at higher 
taxonomic level. Their states for each species 
are shown in the Appendix, and summarized 
for each higher taxon in Table 2. 


> x 
So 
ie 


FIG. 23. Schematic diagrams of 6 types of cartilage tissue. A: Type 1; B: Type 2; C: Type 3; D: Type 


4; Е: Type 5; Е: Туре 6. (See text.) 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 38 


(1) Numbers of Odontophoral Cartilages: In 
gastropods, the number of cartilages varies 
from a single cartilage to five pairs, or are 
lacking. Most commonly, there are one or 
two pairs. When two pairs are present, two 
different types can be recognized — anterior 
and anterolateral pairs of the Acmaeoidea 
(Patellogastropoda) and anterior and pos- 
terior pairs in Vetigastropoda. In the 
Patelloida (Patellogastropoda), there are 
three or more pairs of cartilages including 
an anterior pair. In the Neritomorpha, the 
number of cartilages is typically five (two 
pairs and an unpaired cartilage). In the 
apogastropod clade, there are one pair, a 
single fused piece or none in the Caeno- 
gastropoda, or the cartilages are replaced 
with muscle and connective tissue in the 
Heterobranchia. 

In the phylum Mollusca, odontophoral 
cartilages or corresponding structures are 
present in the “Aplacophora”, Monoplaco- 
phora, Polyplacophora, Scaphopoda and 
Gastropoda, whereas they are lacking in the 
Cephalopoda and Bivalvia (Salvini-Plawen, 
1988; Ponder & Lindberg, 1997: 146; Mes- 
senger & Young, 1999; Table 2). The num- 
ber of cartilages or cartilage-like structures 
is a single pair in aplacophorans (Salvini- 
Plawen, 1988; Scheltema et al., 1994) and 
Scaphopoda (Morton, 1959; Shimek & 
Steiner: 1997: 39—42, fig. 35), one or two 
pairs in Monoplacophora (Lemche & 
Wingstrand, 1959; Wingstrand, 1985: fig. 15; 
Schaefer & Haszprunar, 1996; Haszprunar 
& Schaefer, 1996, 1997), and two pairs in 
Polyplacophora (Wingstrand, 1985: fig. 15; 
Salvini-Plawen, 1988). The number in these 
outgroups suggests that one or two 
odontophoral cartilages or comparable struc- 
tures have occurred in an ancestral group of 
Mollusca, conserved in various molluscan 
classes and subsequently increased or de- 
creased within various groups of gastropods. 


(2) Histology: The tissue of gastropod radu- 
lar-supporting structures can be categorized 
into six types based on the following criteria 
of (a) the presence or absence of the carti- 
lage cells surrounded by the extracellular 
matrix; (b) a relative thickness of the extra- 
cellular matrix around cartilage cells; (c) the 
relative size of the nuclei compared to the 
cartilage cells; and (d) the positions of the 
nuclei within the cartilage cells. 

Type 1: The cartilage matrix is relatively thick; 
compared to other types of tissue, nuclei are 


small and tend to attach to the matrix at the 
edge of the cartilage cells (Fig. 23A). The 
size of nuclei is approximately 7 to 10% of 
cell length. This type is found in Patello- 
gastropoda, Vetigastropoda, and Neriti- 
morpha (Figs. 2-4, 5C—F). 

Type 2: The cartilage is sectored by thick par- 
allel sheets of matrix connected vertically by 
thin layers of matrix; the nuclei are relatively 
large in size and occur in various positions 
within the cartilage cells (Fig. 23B). The size 
of nuclei is around 14% of cell length. This 
type is observed in Cocculinidae (Fig. 5) and 
Lepetelloidea (Haszprunar, 1988c; Hasz- 
prunar & McLean, 1996). 

Type 3: The cartilage matrix is thin and ob- 
scure in outline; nuclei are relatively large 
and can be located in various part of the 
cartilage cells (Fig. 23C). The size nuclei is 
approximately 9-11% of cell length. This type 
is frequently observed in taenioglossate 
Caenogastropoda (e.g., Berthold, 1991; 
Figs. 6, 7, 9-12). 

Type 4: The cartilage matrix is markedly thick; 
extracellular matrix substance stained red by 
Heidenhain’s Azan method extensively fills 
spaces between cartilage matricies; nuclei 
are small relative to cartilage cells and vari- 
ously located within cartilage cells (Fig. 23D). 
The size of nuclei is about 5% of cell length. 
We have only observed this type in the 
caenogastropod family Cypraeidae (Fig. 8). 

Type 5: Cartilage matrix is not as thick as type 
1, but sharply bordered; nuclei are relatively 
large and tend to be located at the center of 
cartilage cells (Fig. 23E). The size of nuclei 
is between 10% and 20% of cell length. This 
type is characteristic of the clade Neogastro- 
poda (Figs. 13-16). 

Type 6: Cartilaginous tissue as defined above 
is absent; the radular bolsters are filled with 
muscular fibers and fibrous connective tis- 
sue which is probably composed of collagen 
fibres (Fig. 23F). This type was observed in 
all the species of Heterobranchia examined 
(Figs. 17-22) regardless of their radular 
types. 

Comparison of outgroups (scaphopods and 
chitons; Fig. 24) suggests that the histology 
of the radular-supporting structures is highly 
diversified in non-gastropod classes. 
“Odontophoral cartilages” in some aplaco- 
phorans are composed mainly of muscle and 
collagen fiber (Scheltema et al., 1994). In 
Polyplacophora, the cartilages are some- 
what similar to type-2 cartilages of Cocculina 
(Fig. 5B). The radular bolster tissue of 


40 


Scaphopoda is similarly composed of muscle 
and fibrous connective tissue (Shimek & 
Steiner, 1997; Fig. 24). In Monoplacophora, 
thin matrix and cartilage cells are observed 
(Haszprunar & Schaefer, 1996). Based on 
recent phylogenetic hypotheses of gastro- 


[3 — um 


KATSUNO & SASAKI 


pods (e.g., Ponder & Lindberg, 1997; Sasaki, 
1998), the primitive state of the cartilage is 
probably type 1, because it is widely shared 
by basal gastropods such as Patello- 
gastropoda, Vetigastropoda, and Neriti- 
morpha. 


FIG. 24. Cross sections of buccal mass of non-gastropod molluscs. А, В: Ischnochiton comptus (Gould, 
1859) (Polyplacophora: Ischnochitonidae). Anterior (A) and middle (B) part of buccal mass; C, D: 
Dentalium octangulatum Donovan, 1804 (Scaphopoda: Dentaliidae). Anterior (C) and posterior (D) 
part of buccal mass; E, F: Episiphon subrectum (Jeffreys, 1883) (Scaphopoda: Gadilinidae). Middle 
(E) and posterior (F) part of buccal mass. 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 41 


In the species we observed, there is no clear 
correlation between the body size and the 
ratio of nuclei size to cell length (e.g., small- 
sized Assiminea vs. other larger caeno- 
gastropods). 


(3) Presence or Absence of an Enclosing 
Membrane: The odontophoral cartilages or 
radular bolsters of molluscs are enclosed by 
a membrane and clearly demarcated from 
surrounding muscles, except some lower 


heterobranchs (Haminoeidae, Philinidae, 
Cylichnidae, Cavoliniidae, and Aplysiidae). 
Since the odontophoral cartilages of non- 
gastropods are also clearly defined, this en- 
veloped type is considered to be primitive in 
Gastropoda. 


(4) Overlapping of Right and Left Main 


Cartilages or Radular Bolsters (Fig. 25): The 
topological condition of the right and left el- 
ements (cartilages and radular bolsters) is 


va 


FIG. 25. Schematic diagrams about the right and left elements of main cartilage or 
radular bolster. A: Ventrally close, connected by double layered ventral approximator 
muscle inserting at ventral/medial side of the main cartilage; B: Ventrally close, con- 
nected by single layered ventral approximator muscle inserting at ventral side of the 
main cartilage or radular bolster; C: Ventrally close, connected by single layered 
ventral approximator muscle inserting at outer lateral side of the main cartilage; D: 
Dorsally close, muscular connection is absent; E: Dorsally close, connected by single 
layered ventral approximator muscle inserting at medial side of the main cartilage. 


42 KATSUNO & SASAKI 


classified into two types; overlapping or not. 
While the former condition is frequently ob- 
served in taenioglossate Caenogastropoda 
(Appendix), the latter is found in other Gas- 
tropoda. 


(5) Closest position of the main cartilage or 
radular bolster in the buccal mass (see Fig. 
25): The main pairs of cartilages or muscu- 
lar radular bolsters may be closest ventrally 
or medially, depending on the taxon. Most 
species observed have ventrally proximal 
cartilages or radular bolsters. However, dor- 
sally proximal cartilages or radular bolsters 
with a convex cross-sectional form are found 
in some taenioglossate caenogastropods 
such as Hipponix (Hipponicidae), Fusitriton 
(Ranellidae), Glossaulax (Naticidae) and 
Strombus (Strombidae) (e.g., Figs. 9C, 11A, 
B, 12). Outgroup comparison suggests that 
ventrally proximal cartilages are found 
throughout Mollusca, except some taenio- 
glossate caenogastropods. These two differ- 
ent types in Caenogastropoda might be 
functionally significant but this meaning can- 
not be clearly elucidated from the limited data 
in this study. 


(6) Insertion of Ventral Approximator Muscle: 

The insertion area of the ventral approxi- 
mator muscle on the odontophoral cartilages 
or radular bolsters is categorized into three 
types, ventral, median, and outer lateral ones 
in gastropods (Fig. 25). The insertion area 
of the ventral approximator muscle is at the 
ventral level of the anterior pair of cartilages 
in Cocculiniformia, Vetigastropoda, Neriti- 
morpha, Neogastropoda, Pulmonata and of 
the ventral layer of Cellana (Nacellidae). The 
ventral approximator muscles in Fusitriton 
(Ranellidae) and the dorsal layer of this 
muscle in Cellana insert at the inner medial 
wall of the cartilages. The insertion of these 
muscles at the outer lateral zone is unique 
to Caenogastropoda except for Neogastro- 
poda. 
The state in outgroups (e.g., Wingstrand, 
1985) indicates that the insertion of the ven- 
tral approximator muscle at the median or 
ventral side of the cartilage is the plesio- 
morphic condition for gastropods. 


Homology 


The homology of different cartilages or radu- 
lar bolsters has been discussed in terms of 


topological relationship, histology, and muscle 
connection (Graham, 1964, 1973; Wingstrand, 
1985; Salvini-Plawen, 1988; Sasaki, 1998; 
Guralnick & Smith, 1999). 

Within gastropods, homology can be estab- 
lished between the anterior cartilages of basal 
gastropods and a single pair or fused carti- 
lage in caenogastropods on the basis of their 
topological correspondence. For the same 
reason, the single pair or single fused carti- 
lage of membrane-enclosed radular bolster in 
Pulmonata can be regarded as a secondarily 
modified homologous structure of an anterior/ 
odontophoral cartilage. These cartilages simi- 
larly support the main part of the odontophore 
and are connected by the ventral approximator 
muscle. The cartilages observed in 
“Aplacophora” (Scheltema et al., 1994) and 
Scaphopoda (Salvini-Plawen, 1988: 363, fig. 
29; Shimek & Steiner, 1997; Fig. 24) seems to 
be the homologue of the anterior cartilages of 
basal gastropods based on topology. 

The homology of polyplacophoran, mono- 
placophoran and patellogastropod radular-sup- 
porting structures has been interpreted in 
different ways. According to Graham (1964) 
and Wingstrand (1985), the anterior cartilage 
of Patellogastropoda is the homologue of the 
polyplacophoran and monoplacophoran radu- 
lar vesicle. However, we question this hypoth- 
esis for two reasons, (1) there is no histological 
evidence to support their interpretation, and (2) 
in Polyplacophora and Monoplacophora, the 
ventral approximator muscle inserting at the 
inner cartilages, not from the radular vesicles 
(Fig. 24). This latter fact, in particular, supports 
the homology between the anterior cartilages 
of patellogastropods and the inner cartilages 
of polyplacophorans and monoplacophorans. 

Graham (1964) argued that the anterior 
cartilages of Patella is derived from the radu- 
lar vesicle by secondary filling of cartilaginous 
tissue, on the ground that the cells of anterior 
cartilage of Patella are notably smaller that 
those of other cartilages. Wingstrand (1985) 
supported Graham’s (1964) hypothesis, and 
also stated that the anterior cartilage of Pa- 
tella is the homologue of the radular vesicle 
and medial (= inner) cartilages. A similar his- 
tology of cartilages was also found in Cellana 
in this study (Fig. 2D), and Graham s (1964) 
and Wingstrand’s (1985) hypothesis seems to 
be supported. However, in the species of 
Acmaeoidea, the histology of the anterior 
cartilages is the same as those of other 
cartilages and also other basal gastropods (e.g., 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 43 


Sasaki et al., 2006b). Therefore, Graham’s 
(1964) and Wingstrand’s (1985) scenario is 
applicable only to the Patellidae and Nacellidae 
and does not apply to patellogastropods as a 
whole. 

The homology of anterolateral cartilages of 
patellogastropods and outer cartilages of 
polyplacophorans and monoplacophorans has 
been suggested by various previous authors 
(Wingstrand, 1985: 62-66; Salvini-Plawen, 
1988: 359-360, Guralnick & Smith, 1999: 184; 
Table 3). This view is supported by the fact 
that the anatomical location of the outer 
cartilages is approximately the same as that 
of the anterolateral cartilage (Figs. 2A, B, 24A, 
B). Similarly, polyplacophoran and mono- 
placophoran inner cartilages seem to be the 
homologue of anterior cartilages of patello- 
gastropods, because they have similar topol- 
ogy in buccal mass and connected by the 
ventral approximator muscle. 


Systematic Implications 


Morphological comparison according to taxo- 
nomic grouping (Appendix, Table 2) revealed 
that most major gastropod groups have char- 
acteristic combinations of character states of 


the odontophoral cartilages or radular bolsters. 

Patellogastropoda: The odontophoral 
cartilages of patellogastropods are mainly 
characterized by (1) more than one pair of 
cartilages (two to five pairs), (2) the presence 
of anterolateral cartilages, and (3) the two lay- 
ers of the ventral approximator muscle (ex- 
cept Bathyacmaea: Sasaki et al., 2006b) 
inserting on the ventral and medial levels of 
the anterior cartilages. 

Vetigastropoda: The majority of veti- 
gastropods have two pairs of cartilages which 
consist of anterior and posterior pairs (Hasz- 
prunar 1987a, 1988b; Sasaki, 1998; this study). 
However, exceptions to this generalization are 
seen in lepetelloideans and “hot-vent” veti- 
gastropods; for example, the anterior and 
anterodorsal cartilages in Choristella (Haszpru- 
nar, 1992) and Bathyphytophilus (Haszprunar 
& McLean 1996), a single pair in Osteopelta 
(Haszprunar, 1988c), Neomphalus (Fretter et 
al., 1981) and Lepetodrilus (Sasaki, 1998), and 
two pairs (anterior and anterodorsal) plus single 
median cartilage in Cocculinella (Haszprunar, 
1988c). Other characters, such as histology 
and the closest point of the anterior pairs, are 
similar to those of patellogastropods and 
Neritimorpha. 


TABLE 3. Statements on patellogastropod cartilages in past studies. (-: no statement). 


Reference Anterior cartilages 


Graham, 1964: 
D. 320 


= mono-/polyplacophoran 
radular vesicle (based on 
histology) 


Anterolateral cartilages 


Posterior cartilages 


Windstrand, 1985: 
pp. 62-66 


= mono/polyplacophoran 
radular vesicle and medial 
cartilage (based on histol- 


ogy, topology) 


= mono/polyplacophoran 
lateral cartilage (based 
on topology) 


= posterior part of poly- 
placophoran lateral 
cartilage (based on 
insertion of some ho- 
mologous muscles) 
Salvini-Plawen, = mono/polyplacophoran = mono/polyplacophoran - 
1988: pp. 359-360 medial cartilage (based on lateral cartilage (based on 
topology) topology) 
plesiomorphic for Mollusca plesiomorphic for Mollusca, - 
(based on position, shape = mono-/polyplacophoran 
and composition) lateral cartilage (based 
on position, shape and 
composition) 


Guralnick & Smith, 
1999: p. 184 


Ad KATSUNO & SASAKI 


Cocculiniformia: This group (in its restricted 
sense) is characterized by having a single pair 
of cartilages with type 2 histology (Haszprunar, 
1987b; Sasaki, 1998; this study). Other char- 
acters are similar to those of other basal gas- 
tropods. 

Neritimorpha: The odontophoral cartilages 
of Neritimorpha are unique in having a single 
median cartilage embedded in the ventral 
approximator muscle between the anterior and 
posterior cartilages (Sasaki, 1998; Kano & 
Kase 2002; Sasaki et al., 2006a). Other char- 
acters are not clearly different from other basal 
gastropods. 

Caenogastropoda: “Lower” Caenogastropo- 
da generally have a single pair of odontophoral 
cartilages (Appendix; Simone, 2001, 2004a, 
b), but the cartilages are fused at anterior end, 
separated, or lost in Neogastropoda (Appen- 
dix; Taylor et al., 1993: 133; Kantor et al., 1997; 
Strong, 2003: character 13). The cartilages are 
missing in part of Conoidea, and this phenom- 
enon is presumably related to the specialized 
venom-injecting mode of feeding with a har- 
poon-like radular tooth. In ptenoglossan fami- 
lies, some parasitic Eulimidae also lacks the 
entire buccal mass due to the reduction of 
mechanical feeding organ (Sasaki et al., 2007: 
see also Waren, 1984). Thus, the loss of the 
cartilages and odontophore can happen in 
some groups with specialized feeding habit. 

Within odotnophore-bearing caenogastro- 
pods, two groups can be recognized histologi- 
cally. (1) In caenogastropods with a 
taenioglossate radula, the cartilages have type 
3 or 4 histology. (2) The members of the clade 
Neogastropoda have type 5 histology defined 
by large nuclei in the cartilage cells surrounded 
by distinct matrix. 

The connection of the ventral approximator 
muscle is always on the ventral side in 
Neogastropoda, but highly variable from ven- 
tral to outer lateral side in taenioglossate 
groups. 

Odontophoral cartilages with distinctive his- 
tology (type 4) are found in Cypraeidae in 
which the cartilage tissue is characterized by 
the existence of an extracellular matrix sub- 
stance that fills large spaces between the car- 
tilage matrices. In addition to Cypraea boivinii 
(Fig. 8), we also confirmed that Cypraea gra- 
cilis japonica has similar histology (personal 
observation), excluding the possibility of an 
abnormality or an exceptional species-specific 
State. It is uncertain whether or not other 


closely related taxa such as Ovulidae have the 
same type of histology. 

Heterobranchia: As well documented in past 
studies (Hyman, 1967: 453; Haszprunar, 
1985a, b, 1988a; Ponder, 1990a, b, 1991; 
Salvini-Plawen, 1988: 332; Luchtel et al., 1997; 
Ponder & Lindberg, 1997: 145), the buccal 
mass of Heterobranchia lacks true cartilages 
and is filled with fibrous connective tissue and 
muscle fibers (Figs. 17-22). As discussed by 
Mackenstedt & Markel (2001: 216-217), this 
type of histology probably allows radular bol- 
sters to change their shape flexibly during 
feeding due to their muscular nature. 

Although our observations on heterobranchs 
are limited, the taxa examined fall into two 
groups. In all but one of the Opisthobranchia 
examined, the radular bolsters are not clearly 
separated from the surrounding tissue (Figs. 
17-18). In contrast, in the pulmonates, the bol- 
sters are clearly demarcated by a membrane 
(Figs. 19-22). The exception in the opistho- 
branchs is the nudibranch Hypselodoris, the 
radular bolsters of which are similar in struc- 
ture to those seen in Pulmonata (Fig. 18E, F). 
The significance of these two types should be 
investigated more extensively throughout the 
heterobranchs in future studies. 

The situation in the lower heterobranchs 
(Heterostropha) is unclear as no non-euthy- 
neuran taxa were investigated in this study. 
However, published studies indicate that these 
taxa too lack either a buccal mass or true 
cartilages (see Appendix for references). 


Evolution of Radula-Supporting Organs in 
Gastropoda 


The evolution of radula-supporting organs 
in Gastropoda can be reconstructed by 
outgroup comparison and tracing the charac- 
ter states on recently published phylogenetic 
trees (Ponder & Lindberg, 1997, for gastro- 
pods; Sasaki, 1998, for basal gastropods; 
Dayrat & Tillier, 2002, for Heterobranchia; 
Strong, 2003 for Caenogastropoda). 

A common gastropod ancestor is inferred to 
possess a combination of the following 
plesiomorphic states in the odontophore: (1) 
two pairs of the odontophoral cartilages, (2) 
unfused right and left elements, (3) type-1 his- 
tology (see above), (4) the ventral insertion of 
the ventral approximator muscle; and (5) clear 
demarcation of odontophoral cartilages from 
the surrounding odontophoral musculature. 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 45 


During the course of evolution in gastropods, 
the number of odontophoral cartilages has 
been greatly diversified. The cartilages have 
increased in part of Patellogastropoda 
(Patellidae and Nacellidae), part of Vetigastro- 
poda (Cocculinellidae), and Neritimorpha, and 
in contrast, fused or lost in Neogastropoda, or 
replaced by muscles and fibrous connective 
tissue in Heterobranchia. In earlier studies, 
Hyman (1967: 241) stated that the presence 
of several radular bolsters (odontophoral 
cartilages) is primitive, and that the reduction 
to one pair due to fusion is a derived condition. 
Meanwhile, Salvini-Plawen (1988: 359-360) 
advocated that higher numbers of cartilages 
in Patellidae and one pair or single fused pair 
in neomphalid or caenogastropod cartilages 
are secondary products. Similarly, Ponder & 
Lindberg (1997: 146) also regarded the in- 
crease in cartilage number as secondary. In 
accordance with these recent studies, the lat- 
ter scenario is better supported in this study. 

Evolutionary changes of radula-supporting 
structures at a histological level have been 
rarely discussed in the past. In this study, primi- 
tive type-1 histology of basal groups is as- 
sumed to have been modified into types 3, 4, 
5 in Caeogastropoda and type 6 in Hetero- 
branchia, respectively. 

The odontophoral cartilages or radular bol- 
sters are enclosed by a sheath of collagen fi- 
bers in all gastropods other than some 
opisthobranchs (Haminoeidae, Philinidae, 
Cylichnidae, Cavoliniidae, and Aplysiidae: 
Figs. 17, 18A-D; Appendix). Thus, ensheathed 
cartilages are regarded as original gastropod 
condition, and the loss of the sheath is 
apomorphic change occurred in part of 
Opisthobranchia. 

The differences in the insertion areas of the 
ventral approximator muscle seem to be re- 
lated to the lateral rotation of the main 
cartilages during feeding. Ventrolateral inser- 
tion allows lateral rotation of the radula (Fretter 
& Graham, 1962; Hyman, 1967; Graham, 
1973; Salvini-Plawen, 1988), but doso- or ven- 
tromedial insertion does not. The former in- 
sertion is regarded as advanced and observed 
frequently in Gastropoda, except for 
taenioglossate caenogastropods, which have 
outer lateral insertion. The insertion of the ven- 
tral approximator muscle might be one factor 
affecting the mode of feeding as Salvini- 
Plawen (1988) stated, but at present we are 
unable to give a detailed account. 


A correlation between morphological types 
of the radula-supporting organs and modes 
of feeding is clearly rejected in most cases. 
For example, various herbivorous gastropods 
belonging to Patellogastropoda, Vetig- 
astropoda, Aplysia and Siphonaria have dis- 
tinctive morphology and/or histology that is 
specific to each higher taxonomic category. 
Similarly, species in the caenogastropod 
Littorinimorpha share similar cartilage mor- 
phology and histology but show highly diverse 
modes of feeding such as grazing in Littorina, 
shell boring in Naticidae, deposit feeding in 
Strombidae, ciliary feeding in Calyptraeidae 
and carnivory in Ranellidae. Thus, there is no 
direct connection between odontophoral car- 
tilage morphology/histology and feeding ecol- 
ogy in gastropods. 

Major habitat selection is also unrelated to 
morphological diversification of radular sup- 
porting organs. Different freshwater or terres- 
trial groups do not fall into the same category 
but share similar structures with phylogeneti- 
cally closer members in marine environments. 
Accordingly, the morphological diversity in 
radula-supporting organs must be a phyloge- 
netically constrained phenomenon. 

In conclusion, the diversification of gastro- 
pod radula-supporting structures is virtually 
controlled by phylogeny regardless of feeding 
ecology and habitat. The functional differen- 
tiation of the odontophore must be considered 
in connection with comparative anatomy of 
buccal musculature and development of dif- 
ferent types of proboscis in future studies. 


ACKNOWLEDGEMENTS 


We deeply thank anonymous reviewers, 
Prof. George M. Davis (George Washington 
University), Dr. Winston F. Ponder (Australian 
Museum), Dr. Amélie H. Schetema (Woods 
Hole Oceanographic Institution), and Prof. 
Kazushige Tanabe (University of Tokyo) for 
substantial improvement of the manuscript. 
We also acknowledge kind assistance in sam- 
pling by Dr. Tatsuo Oji (University of Tokyo), 
Dr. Takao Ubukata (Shizuoka University), Dr. 
Kotaro Tsuchiya (Tokyo University of Marine 
Science and Technology), and Prof. Yoshihisa 
Shirayama (Seto Marine Biological Station, 
Kyoto University). This study was supported 
by a Grant-in-Aid from Japan Society of the 
Promotion of Science (No. 18770063). 


46 KATSUNO & SASAKI 


LITERATURE CITED 


BARKER, G. M., 2001, Gastropods on land: phy- 
logeny, diversity and adaptive morphology. Pp. 
1-146, in: G М. BARKER, ed., The biology of ter- 
restrial molluscs. New York, CABI Publishing. 
xiv + 558 pp. 

BERTHOLD, T., 1991, Vergleichende Anatomie, 
Phylogenie und historische Biogeographie der 
Ampullariidae (Mollusca, Gastropoda). Ab- 
handlungen des Naturwissenschaftlichen 
Vereins in Hamburg, 29: 1-256. 

BOUCHET, Р. 4 J. P. ROCROI, 2005, Classifi- 
cation and nomenclator of gastropod families. 
Malacologia, 47(1-2): 1-397. 

COLE, А. С. & В. К. HALL, 2004, Cartilage is a 
metazoan tissue; integrating data from nonver- 
tebrate sources. Acta Zoologica (Stockholm), 
85: 69-80. 

DAVIS, G. M., 1967, The systematic relationship 
of Pomatiopsis lapidaria and Oncomelania 
hupensis formosana (Prosobranchia: Hydrobii- 
dae). Malacologia, 6: 1-143. 

DAYRAT, B. & S. TILLIER, 2002, Evolutionary 
relationships of euthyneuran gastropods (Mol- 
lusca): A cladistic re-evaluation of morphologi- 
cal characters. Zoological Journal of the 
Linnean Society, 135: 403-470. 

FRETTER, V., 1988, New archaeogastropod lim- 
pets from hydrothermal vents, superfamily 
Lepetodrilacea. Il. Anatomy. Phylosophical 
Transactions of the Royal Society of London, 
(B), 319: 33-82. 

FRETTER, V., 1989, The anatomy of some new 
archaeogastropod limpets (superfamily 
Peltospiracea) from hydrothermal vents. Jour- 
nal of Zoology, 218: 123-169. 

FRETTER, V. & А. GRAHAM, 1962, British proso- 
branch molluscs. Their functional anatomy and 
ecology. London, The Royal Society. 755 pp. 

FRETTER, V., A. GRAHAM & J. H. MCLEAN, 
1981, The anatomy of the Galapagos Rift lim- 
pet, Neomphalus fretterae. Malacologia, 21(1- 
2) 391301. 

GRAHAM, À., 1964, The functional anatomy of 
the buccal mass ofthe limpet (Patella vulgata). 
Proceeding of the Zoological Society of Lon- 
don, 143: 301-329. 

GRAHAM, A., 1965, The buccal mass of ianthinid 
prosobranchs. Proceedings of the Malacologi- 
cal Society of London, 36: 323-338. 

GRAHAM, A., 1973, The anatomical basis of 
function in the buccal mass of prosobranch and 
amphineuran molluscsc. Journal of Zoology, 
169: 317-348. 

GURALNICK, R. & K. SMITH, 1999, Historical 
and biomechanical analysis of integration and 
dissociation in molluscan feeding, with special 
emphasis on the true limpets (Patello- 
gastropoda: Gastropoda). Journal of Morphol- 
ogy, 241: 175-195. 

HASZPRUNAR, С., 1985a, Zur Anatomie und 
systematischen Stellung der Architectonicidae 
(Mollusca, Allogastropoda). Zoologica Scripta, 
14(1): 25-43. 


HASZPRUNAR, G., 1985b, On the anatomy and 
systematic position of the Mathildidae (Mol- 
lusca, Allogastropoda). Zoologica Scripta, 14: 
201-213. 

HASZPRUNAR, G., 1987a, The anatomy of 
Addisonia (Mollusca, Gastropoda). Zoo- 
morphology, 106: 269-278. 

HASZPRUNAR, G., 1987b, Anatomy and affini- 
ties of cocculinid limpets (Mollusca, Archaeo- 
gastropoda). Zoologica Scripta, 16(4): 305-324. 

HASZPRUNAR, G., 1988a, On the origin and 
evolution of major gastropod groups, with spe- 
cial reference to the Streptoneura. Journal of 
Molluscan Studies, 54: 367-441. 

HASZPRUNAR, G., 1988b, Anatomy and affini- 
ties of pseudococculinid limpets (Mollusca, 
Archaeogastropoda). Zoologica Scripta, 17(2): 
161-179. 

HASZPRUNAR, G., 1988c, Anatomy and rela- 
tionships of the bone-feeding limpets, 
Cocculinella minutissima (Smith) and Osteo- 
pelta mirabilis Marshall (Archaeogastropoda). 
Journal of Molluscan Studies, 54: 1-20. 

HASZPRUNAR, G., 1989a, The anatomy of 
Melanodrymia aurantiaca Hickman, a coiled 
archaeogastropod from the east pacific hydro- 
thermal vents (Mollusca, Gastropoda). Acta 
Zoologica (Stockholm), 70(3): 175-186. 

HASZPRUNAR, G., 1989b, New slit-limpets 
(Scissurellacea and Fissurellacea) from hydro- 
thermal vents. Part 2. Anatomy and relation- 
ships. Contributions in Science, Natural History 
Museum of Los Angeles County, 408: 1-17. 

HASZPRUNAR, G., 1992, On the anatomy and 
relationships of the Choristellidae (Archaeo- 
gastropoda: Lepetelloidea). The Veliger, 35(4): 
295-307. 

HASZPRUNAR, G. & J. H. MCLEAN, 1996, 
Anatomy and systematics of bathyphytophilid 
limpets (Mollusca, Archaeogastropoda) from 
the northeastern pacific. Zoologica Scripta, 
25(1): 35-49. 

HASZPRUNAR, G. & K. SCHAEFER, 1996, 
Anatomy and phylogenetic significance of 
Micropilina arntzi (Mollusca, Monoplacophora, 
Micropilinidae Fam. Nov.). Acta Zoologica, 77 
(4): 315-334. 

HASZPRUNAR, С. & К. SCHAEFER, 1997, 
Monoplacophora. Pp. 415-457, in: F. W. 
HARRISON & A. J. KOHN, eds., Microscopic 
anatomy of invertebrates. 6B: Mollusca Il. New 
York, Wiley-Liss. xvi + pp. 415-828. 

HUBENDICK, B., 1978, Systematic and com- 
parative morphology of the Basommatophora. 
Pp. 1-47, in: V. FRETTER & J. PEAKE, eds., Pul- 
monates, Vol. 2A. Systematics, evolution and 
ecology. London, Academic Press. xi + 540 pp. 

HYMAN, L., 1967, Mollusca |. The Invertebrates: 
Vol. VI. New York. McGraw-Hill. 792 pp. 

KANO, Y. & T. KASE, 2002, Anatomy and sys- 
tematics of the submarine-cave gastropod 
Pisulina (Neritopsina: Neritliidae). Journal of 
Molluscan Studies, 68: 365-384. 

KANTOR, Y. I., 1991, On the morphology and 
relationships of some oliviform gastropods. 
Ruthenica, 1: 17-52. 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 47 


KANTOR, Y. I. & M. С. HARASEWYCH, 1992, 
Morphology of the digestive system of 
Volutomitra alaskana Dall, 1902 (Gastropoda, 
Pectinibranchia, Volutomitridae), with notes on 
a possible mechanism of feeding. Ruthenica, 
2: 45-53. 

KANTOR, У. 1. & А. |. MEDINSKAYA, 1991, Mor- 
phology and feeding of Mitrella burchardi (Gas- 
tropoda: Columbellidae). Asian Marine Biology, 
8: 25-33. 

KANTOR, У. 1., А. MEDINSKAYA & J. D. TAY- 
LOR, 1997, Foregut anatomy and relationships 
of the Crassispirinae (Gastropoda, Conoidea). 
Bulletin of the Natural History Museum, Lon- 
don (Zoology), 63: 55-92. 

KANTOR, Y. I. & J. D. TAYLOR, 1994, The fo- 
regut anatomy of Strictispira paxillus (Reeve, 
1845) (Conoidea: Strictispiridae). Journal of 
Molluscan Studies, 60: 343-346. 

LEMCHE, Н. & К. С. WINGSTRAND, 1959, The 
anatomy of Neopilina galatheae Lemche, 1957 
(Mollusca, Tryblidiacea). Galathea Report, 3: 
9-72. 

ГОСНТЕЕ О... А: Е. MARTIN, I. DETRUP- 
OLSEN & H. H. BOER, 1997, Gastropoda: 
Pulmonata. Pp. 459-718, in: F. W. HARRISON & 
A. J. KOHN, eds., Microscopic anatomy of in- 
vertebrates, 6B: Mollusca Il. New York, Wiley- 
Liss. xvi + рр. 415-828. _ 

MACKENSTEDT, U. & K. MARKEL, 2001, Radu- 
lar structure and function. Pp. 213-236, in: С. 
M. BARKER, ed., The biology of terrestrial mol- 
luscs. New York, CABI Publishing. xiv + 558 


Pp. 

MESSENGER, J. В. & J. Z. YOUNG, 1999, The 
radular apparatus of cephalopods. Philosophi- 
cal Transactions of the Royal Society of Lon- 
don, (B), 354: 161-182. 

MORTON, J. E., 1959, The habits and feeding 
organs of Dentalium entalis. Journal of the 
Marine Biological Association of the United 
Kingdom, 38: 225-238. 

PONDER, W. F., 1970, Some aspects of the 
morphology of four species of the neo- 
gastropod family Marginellidae with a discus- 
sion on the evolution of the toxoglossan poison 
gland. Journal of Malacological Society of Aus- 
tralia, 2: 55-81. 

PONDER, W. F., 1972, The morphology of some 
mitriform gastropods with special reference to 
their alimentary and reproductive systems 
(Neogastropoda). Malacologia, 11: 295-342. 

PONDER, W. F., 1986, Glacidorbidae 
(Glacidorbacea: Basommatophora), a new 
family and superfamily of operculate freshwa- 
ter gastropods. Zoological Journal of the Lin- 
nean Society, 87: 53-83. 

PONDER, W. F., 1987, The anatomy and rela- 
tionships of the pyramidelacean limpet 
Amathina tricarinata (Mollusca: Gastropoda). 
Asian Marine Biology, 4: 1-34. 

PONDER, W. F., 1990a, The anatomy and rela- 
tionships of the Orbitestellidae (Gastropoda: 
Heterobranchia). Journal of Molluscan Stud- 
ies, 56: 515-532. 


PONDER, W. F., 1990b, The anatomy and rela- 
tionships of a marine valvatoidean (Gas- 
tropoda: Heterobranchia). Journal of Molluscan 
Studies, 56: 533-555. 

PONDER, W. F., 1991, Marine valvatoidean gas- 
tropods — implications for early heterobranch 
phylogeny. Journal of Molluscan Studies, 57: 
21-32. 


PONDER, W. F. & D. R. LINDBERG, 1997, To- 
wards a phylogeny of gastropod molluscs: an 
analysis using morphological characters. Zoo- 
logical Journal of the Linnean Society, 119: 83- 
269. 

RUTHENSTEINER, B., 2006, Redescription and 
3D morphology of Williamia gussonii (Gas- 
tropoda: Siphonariidae). Journal of Molluscan 
Studies, 72: 327-336. 

SALVINI-PLAWEN, L. V., 1988, The structure and 
function of molluscan digestive systems. Pp. 
301-380, in: E.R. TRUEMAN & М. В. CLARKE, eds., 
The Mollusca. Vol. 2: Form and function. San 
Diego, California, Academic Press. xxviii + 504 


рр. 

SALVINI-PLAWEN, L. V. & С. STEINER, 1996, 
Synapomorphies and plesiomorphies in higher 
classification of Mollusca. Pp. 29-52, in: J. D. 
TAYLOR, ed., Origin and evolutionary radiation 
of the Mollusca. Oxford, U.K., Oxford Univer- 
sity Press. xiv + 392 pp. 

SASAKI, T., 1998, Comparative anatomy and phy- 
logeny of the recent Archaeogastropoda (Mol- 
lusca: Gastropoda). The University Museum, the 
University of Tokyo, Bulletin, 38: 1-223. 

SASAKI, T., К. MURO & M. KOMATSU, 2007, 
Anatomy and ecology of the shell-less en- 
doparasitic gastropod, Asterophila japonica 
Randall and Heath, 1912 (Mollusca: 
Eulimidae). Zoological Science, 24: 700-713. 

SASAKI, T., Т. OKUTANI & К. FUJIKURA, 2006a, 
Anatomy of Shinkailepas myojinensis Sasaki, 
Okutani, and Fujikura, 2003 (Gastropoda: 
Neritopsina). Malacologia, 48(1-2): 1-26. 

SASAKI, Т., T. OKUTANI & К. FUJIKURA, 2006b, 
Anatomy of Bathyacmaea secunda Sasaki, 
Okutani, and Fujikura, 1993 (Patello- 
gastropoda: Acmaeidae). Journal of Molluscan 
Studies, 72: 295-309. 

SCHAEFER, K. & G. HASZPRUNAR, 1996, 
Anatomy of Laevipilina antarctica, a mono- 
placophoran limpet (Mollusca) from Antarctic 
waters. Acta Zoologica, 77: 295-314. 

SCHELTEMA, A. H., M. TSCHERKASSKY & A. 
M. KUZIRIAN, 1994, Aplacophora. Pp. 13-54, 
in: F. W. HARRISON & А. J. KOHN, eds., Micro- 
scopic Anatomy of Invertebrates. 5: Mollusca 
I. New York, Wiley-Liss. xiv + 390 pp. 

SHIMEK, R. L. & G STEINER, 1997, Scapho- 
рода. Pp. 719-781, in: Е. W. HARRISON & A. J. 
KOHN, eds., Microscopic anatomy of inverte- 
brates, 6B: Mollusca Il. New York, Wiley-Liss. 
xvi + pp. 415-828. 

SIMONE, L. R. L., 2001, Phylogenetic analysis 
of Cerithioidea (Mollusca, Caenogastropoda) 
based on comparative morphology. Arquivos 
de Zoologia, 36(2): 147-263. 


48 KATSUNO & SASAKI 


SIMONE, L. R. L., 2004a, Morphology and phy- 
logeny of the Cypraeoidea (Mollusca, 
Caenogastropoda). Rio de Janeiro, Papel Vir- 
tual Editora. 184 pp. 

SIMONE, L. R. L., 2004b, Comparative morphol- 
ogy and phylogeny of representatives of the 
superfamlies of architenioglossans and the 
Annulariidae (Mollusca, Caenogastropoda). 
Arquivos do Museu Nacional, Rio de Janeiro, 
62: 384-504. 

STRONG, E. E., 2003, Refining molluscan char- 
acters: morphology, character coding and a 
phylogeny of the Caenogastropoda. Zoologi- 
cal Journal of the Linnean Society, 137: 447- 
554. 

TAYLOR, J. D., Y. I. KANTOR & А. Y. SYSOEV, 
1993, Foregut anatomy, feeding mechanisms, 
relationships and classification of the Conoidea 
(= Toxoglossa) (Gastropoda). Bulletin of the 
Natural History Museum, London (Zoology), 
59: 125-170. 

TAYLOR, J. D. & J. A. MILLER, 1990, Anew type 
of gastropod proboscis: the foregut of Hastula 


bacillus (Gastropoda: Terebridae). Journal of 
Zoology, 220: 603-617. 

VOLTZOW, J., 1994, Gastropoda: Proso- 
branchia. Pp. 111-252, in: Е. W. HARRISON & А. 
J. KOHN, eds., Microscopic anatomy of inverte- 
brates, Vol. 5: Mollusca |. New York, Wiley-Liss. 
xiv + 390 pp. 

WAREN, A., 1984, A generic revision of the fam- 
ily Eulimidae (Gastropoda, Prosobranchia). 
Journal of Molluscan Studies, Supplement, 13: 
1-96. 

WAREN, A. & P. BOUCHET, 2001, Gastropoda 
and Monoplacophora from hydrothermal vents 
and seeps; new taxa and records. The Veliger, 
44: 116-231. 

WAREN, A. & W. F. PONDER, 1991, New spe- 
cies, anatomy and systematic position of the 
hydrothermal vent and hydrocarbon seep gas- 
tropod family Provannidae fam. n. (Caeno- 
gastropoda). Zoologica Scripta, 20: 27-56. 

WINGSTRAND, K. G., 1985, On the anatomy and 
relationships of Recent Monoplacophora. 
Galathea Report, 16: 7-94. 


Revised ms. accepted 5 September 2007 


49 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 


(senunuos) 


(6 ‘v-z "SB :9661) 


s|¡99 aBe]1ieo 


Jaj9eyos $ Jeunidzsey 2 7 juesge Jussaud pue хеш и! Z шие виа Seplul|Idosol|/\ 
(6 ‘с "SB :9661) eopoJeJue 
JeunidzseH Y Jejoeyos a ueIpau juasqe juasaJd ¿ zZ eujidinae7 эерииаоэм 
иерэш seayjejeb 
(5861) рчелзбилл  лэчищедиэл ¿Jenuan juasqe Juosoid 2 (ono ‘5! y euljidoan эерииаоэм 
иерэш Jauul 
(5861) рчелзби\л nenusA едиэл juasqe Juosaid & (ono ‘5! y била ешэл эерщиэд 
eolydosejdouoA 
иерэш $99 aBejnueo snjjese 
(5861) puensBuim  лэчищелдиэл ¿Jenuan juasge Jussaud pue хеш ul] (ono ‘ol) 7 sninajdopida7 sepiuoyyoojda7 
(а $195 aBejnueo snjduioo 
—YVpyz ‘s6i4) Apnis SIUL uelpew Jeuul едиэл Juosqe Juosaid pue хщеш ul] (ano ‘5! y UOJYIOUYIOS] SeplUOYYIOUYOS] 
черэш 
(5861) puensBuim лэчищедиэл 5едиэл juasqe juasa.d ¿ (ono ‘91) y ‘ds xeydoziy2S Sepluoyyoouyos] 
e1oudo92e|do{Âjo4 
SIOQI} 
(2661) Je ja euwayeyos ¿ ¿ juesge ¿ UePe]|09 pue ajosnuu с sn3snqol sndopnas  Sepuossoyuur] 
s|99 эх|-эбецлеэ 
UM $494 Juenf 
(1661) je 19 euwayayos леэрип едзиэл juesge juasald ueBe]lo9 pue ajosnuu с виэшореюэфрэн  *SepielyjoJwuis 
елоцаоэе 
ээцэлээч ajosnu SJ8]S|0q SJ9]S|0Q JE] J3]S|0Q ÁBOJO1SIH s19]S/0q лепрел зэюэа< exe | JayBiH 
Joyewixoidde  Jejnpe. -npe Jo $эбе|  Jejnpel JO зэбенеэ 
|¡e1Juan Jo saPejnieo = -meourew Jo эбещлеэ Jo JaquinN 
jo uolesu; меш jo uo] ye; риезубиы Hulsojous 


-1504 159500) JO биааенэло euelquen 


‘(L00Z) зэчэпоя $ usen, зиэзае зэбешиеэ :- ‘шецээип 
:¿ Bees Jajno :90 ‘эбешаеэ Jouul :91 ‘эбешиеэ |езлоро.эзие :эре ‘эбепиеэ черэш :эш ‘эбешаеэ jesiopoiaisod :opd ‘sBele9 Jola3sod :od ‘abel 
-J89 |елэ}е| зомззие :oje ‘obeyed |едиэл :эл ‘эбелеэ Jolajue :эе ‘зэзп|ош Jeyjo pue spodo.]se6 и! s1a]sjoq Jejnpe. Jo saBejnieo Jo saje]s лэоелец 


XIGNAddV 


KATSUNO & SASAKI 


50 


(senunuoo) 


ери/еа 
(981 ‘бу :8661) Meses ¿uelpau Jauul едиэл é juesold a (je '98) y e/n399709AIN эерэешэ\ 
(9-98 `$би epun2es 
'q900Z) ‘Je 19 Meses едиэл едиэл juasqe Juoseld | odA (je '98) y eoewaefiyeg SEPISEWOY 
(q-e21 едиеэл з/ешиано 
"SB :866L) Meses  /Ueipelu JeuuI едиэл juasqe Jussaud & (oye ‘oe) y EJUOPOUNIAA эерэешюо\ 
[EUSA 
(АР, "By :8661) Meses /ueipeuwu Jauul jenusan Juosqe juosaid Ip (se 9e) y вии вэаавиит эерцэаэ] 
(эра 
(е/ ‘Bu :8661.) Meses À é juasqe Jussaud ¿ ‘od ‘oye ‘OA ‘эе) QOL  ELUN810] вие]эЭ Sepl||SOeN 
едиэл (эра 
(2 ‘VL :s614) Apnis SIUL /иерэшаэиш едиэл Juosqe JuasaJud | adÁ] ‘od ‘oye ‘an 98) OL eJeıb eue/[81 эернээем 
ев bi? едиэл (эра 
‘с “SB :ÿ96L) Weyelo /черашаэци Oh juasqe juasa.d ¿ ‘od ‘oye ‘On '98) OL влеб/пл ejeed эериэ}еа 
(5 езопха 
—ez ‘SB :8661) Meses ¿uelpau Jauul [EUSA juesge Ju9sald a (od ‘oye ‘oe) y 2/1S2/81n9S 3€pI|1918 4d 
epodonseßojlsjed 
SJOQI 
(4 леюзпш pue эп$$1 wnjoaigns 
—Jyz ‘s6i4) Apnis sy [EUSA jenusA Jussqe juasqe = элцоэчциоэ зполац С uoydisidz эершииреэ 
SOI] 
(a Jejosnw pue эп$$1 ште/бие]о 
—Э уе ‘614) Apnis sy ¡esop ¡esJop juasqe JuUasald — элцоэчциоэ SNO.IQU С шпиеиэа эершаиэа 
(2661) éSII89 
J3UI9IS Y Yawıys едиэл едиэл Juosqe Jussaud pally-piny 9618] с  sueuage ejipes gepIlıpes) 
(1661) 
JOUIS]S Y YSWIUS [EUSA [EUSA Jussqe Jussaud é с  sn1991 зпраецыу эерраечч 
epodoydeos 
a9U9J9/9y ajosnu SJ9]S|0Q SJ3]S|0Q JE] J9]S|0Q ÁBOJOISIH sJ8]S|0q Jenpel зэюэас exe | J9SyBiH 
Joyewixoidde Jepnpel -Npe Jo sabe] Jenpe, JO soBeliue9 
¡e1Juen Jo зэбещлеэ -meourew Jo obeyed Jo JOQUINN 
jo чоцлэзч|  UIEU JO UOI Je] рчезчбы Bulsojous 
-ISOd 1$9$0|9 Jo Buiddejlano eueiquen 
(репициоэ) 


51 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 


(senunuoo) 


(vr By snj99/9p 
:d6861) JeunidzseH ¿Jenuen езчэл juasqe juasaJd | adh} (od ‘oe) y snjoasosdÁlO —„эери.ро}э4э7 
snsojnisnd 
(886L) 191914 é ¿ E juasaJd ¿  SJled z 1See] Je snppojada7 aepiupojada7 
(р-959 
"sb :8661) Meses едиэл [EUSA juesge juasaJd é с xXnusnjupojeda7  eepl|ipoyede] 
(y By juaspnuy эер!и! 
:4886L) JeunidzseH E é E juasaJd ¿ (od ‘oe) y eoijdimyduwy = -пээоэорпэза 
(e “By SISUBJONZBUBA эершн 
:49961) seunsdzseyH [EUSA é juesge 1uasa.d E (od ‘oe) y eissAgejuny = -пээоэорпэза 
(vr ‘By 
‘:2/96L) Jeunidzse}J é é Py Juosaid 2 (od ‘oe) y S//218]2] ешозрру эер|э1э9э7 
(91 “By $/!делии 
98861) JeunsdzseH 2. ¿ juasqe Juosa.d ¿ Z eyjadoajsC sepIıyadosIso 
(er ‘LL ‘2 ‘6 “sby (ou) & (oui EUIISSHNUIU 
:0886L) Jeunidzse}J едиэл езиэл éjuesge juasald ‘(ope) ¿ ‘(эе) г adAj эби$ ‘ope ‘эе) © ejjauln9909  2EPIJOUINN9909 
(с e ‘sby seuewyaly 
:Z66L) Jeunidzsey й & juesge Juosa:d ¿e edÁ] (ope ‘эе) y e//AJSUOYD SEPI||OISUOUD 
(21-91 ‘S64 :9661) sisuebalp Sep 
ues TOW Y Jeunidzsey & езичэл éjuesqe juasald (ope) ‘(эе) z adh} (эре '9e) y snyiydojAydAyjeg = -1ydo\AyudAujeg 
sııyenbe 
(p-91€ JOJOIISISNIP 
‘SIL :28661) Meses едчэл езчэл juasqe Ju9sa.Jd 2 (od ‘oe) y sninojns эерпонен 
sııyenbe 
(£ JOJOIISAISAIP 
‘a, ‘s6i4) Apnis sy] JeJJUSA езчэл juasqe jJu3sa.Jd | adÁ1 (od ‘эе) y SNONECH эерпонен 
(94-295 
‘SBU :8661) Meses едиэл езиэл juasqe jJuasaJd ¿ (od '9e) p  sisuauls sninas Ssepi|jaunssi- 
ерододзебцэл 
ээцэ.ээч SpasnW Si9]S|0Q SJ9]S|0Q 8] J9]S|0Q ÁBOJO1SIH Si9]S|0q Je]npeJ sardads exe | JeUBIH 
Joyewixoidde Jeinpe\i -прел Jo sebe| Jepnpei Jo sebeliue9 
едчэл Josebeyeo -meourew Jo эбещаеэ Jo JOQUINN 
jo UOILISSU]  UIEUI JO UOI ys] pue зчбы Buisojous 


-ISod jsesojg JO Buiddeusno эчелашэи 


(репициоэ) 


(senunuos) 


(4-96 ‘94 'sBiy) 


Apnjs $14} ‘(р-96/ (ou) $ adh} 
`5би :8661) Meses [EUSA елиэл juasqe juasald ‘(ad ‘oe) | edA] (our ad" Se) e e//191Q/8 ешэм SEPNMON 
ечалошпиэм 
(£ epidsiy 
‘BU :9/861) seunsdzseH едчэл езиэл juasqe juasaJd ¿ Z e/bAd093909 BEPIUINNII09 
esıuoddiu 
(p69 ‘Bu :8661) Meses елчэл едиэл juasqe JU3SsaJd é С eCUIINDIOD BEPIUININII0I 
esıuodel 
(g-ys ‘sbi4) Apnis SIUL езиэл езиэл juasqe jJuasa.Jd с adh} Z “yO ‘ds eulna90) 9EPIUNNII0I 
EILUIOJIUIIN9909 
(q-ees 194Y98! 
г. ‘SB :8661) Meses едиэл [EUSA juasqe Juosa.d a (od '98) y CWOASOIO[YD эерщэо. | 
— esnjuod 
(09) (y 'B14) Apnıs $141 ¡e1Juan елиэл juesge Juosa.d | adÁ1 (od '9e) y ое! виороиои| oepIU901L 
< 
9%) !youAeq 
où (poz ‘By :8661) Meses едиэл елиэл juasqe juasa.Jd & (od '98) y sny90.10PIyy эершешоо.пэ|а 
O (9 ‘G-r Zz sb esenuelne 
5 :6861) Jeunidzsey é é é Juesa1d ¿e adh} лед | 1ses] Je виилроие!э// эершазоэа 
р COMJUBDUOD 
< (G—p ‘SB :6861) 191914 [EUSA едиэл juasqe ¿Jussald é 2 eyedoyouAyH эершазо\эа 
(Sy 921901] 
“SHU :1861) Je 19 19N914 ¿ é é]uesge juasaJd ¿ Z snjeydwoey vepijeydwoaN 
(9-Y| ‘su зааипа 
:4686L) seunudzsey ¿Jenuean [EUSA juesge Juasaid | эал} (od ‘oe) y SIJDUIDOUW9 | SEPIUOZIIINS 
snyeulbsewa 
(21 “By :8861) 187214 é ¿ ¿ jueseid à 3164 с See] je sydajobio5 „эернирозэаел 
(V6 ‘9 ‘ve ‘sby эвивиеш 
:9686 1) Jeunsdzsey ¿Jenuan [EUSA juesge Juosald | adÁ1 (od ‘эе) y enuniopnasg  ,sepijmpojeda7 
39U9J9/9y aJosnu sJ9]S|0Q SJ3]S|0Q JE] J9]s|0q ÁBOJO1SIH SJ9]S|0q лепрел sennads exe | JayBin 
Joyewixoidde лепрел -Npe. Jo see] Jejnpes JO зэбееэ 
езиэл Jo зэбецлеэ wed меш JO 288/1189 JO JOQUINN 
jo uolesu]  UIEW JO UON 19] pue зубы Bulsojoue 
-Isod 159$09 jo биааемэло эчелашэи 
N 
LO (репициоэ) 


95 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 


(зэпипциоэ) 


(d-v6 
‘3, ‘s6i4) Apnis SIUL  /eJaje] tajo едиэл Juosaid Juasaid с odA] с Eejnaınaıg CULIO}IT 38pIULIO1)17] 
(8 Oz :sB14) Apnis sy | [EUSA [EUSA juasqe juasaJd y adÁ] с  HUINIOQ eaeidAD эерэела^э 
(za) 
‘4, `$61-1) Apnis siuj [2192] Jano езчэл juasqe Juosaid с edA] с хлио епраэл  eepisendAe9 
(q-vz `$614) Apmis siuj  /eJaje] Jano едиэл juesald juesald с edA] с ибишипэ веиещея эершепеа 
(8v2 
‘BY :1661) ployuag @ é é jueseld é 2 вади eses Bepiuejnduy 
(бус ‘/ус snapnouejeq 
‘SOU :1661) Рючыэа  ¡eJejej 180 é juesaid juesald & 7 snwodoyy  sepiuejndwy 
(182 ‘092 SHOUENUIO9 
‘SIL :1661) рючыэа & едичэл é jueseld с edA] Z esueyy — эершетпашу 
(9-99 eJ98] SISUBUIYO 
‘ар ‘s6i4) Арп}$ $41 [21972] Jajno едиэл }иэ$э.а juasad © edA] с euipnjedoßuedig ABPHECIAIA 
IS]O419Y 
(9—\9 ‘sbi4) Apnis SIUL  /8/aje] это едичэл }иэ$э.а Juosa.d с edA] 2 snioydoj3/9) svepl1oydojoAQg 
(OLL ‘VOL ‘V6 “sb SIJIQEUEA 
:1661) лэрчоа Y чэлем é é у juaesaid с edA] Z 2uuenrold эершиело.а 
epodonseßousen 
ajesso¡Boluae | 
(q-vg ‘sb sisuaulloAw эер 
89007) ‘Je 19 Meses едчэл едиэл juasqe Juosa.d | odA} (ou ‘od ‘oe) в sedajieyulyS -ıpedsjoseusyd 
(q-ee8 в/ецота эер 
`5бу :8661) Meses едиэл едиэл juesge juosold é (Эш ‘od ‘эе) с ejadajeuuly -ıpedsjoseusyd 
D—068 esıuodel 
“SOU :866L) Meses [EUSA езиэл juasqe juasa.d é (ou ‘od ‘эе) y euiewapjey эерщиэнэн 
(39 eueiswepe 
‘By :Z00Z) SSEN Y OUBY едиел é juesqe é é (dW ‘od ‘oe) в BUIINSId MEN 
3249.99 y a19snu SJ9]S|0Q SJ91S|OQ Je} Ja\sjog ÁBOJOISIH SJ9]S|0q Je¡npe, saldads exe | JaybiH 
Joyewuixoidde Jepnpel -пре/ Jo see] Jejnpes Jo зэбееэ 
ENTES JO sebeyieo -weo меш Jo эбещлеэ JO JOQUINN 
jo UOIHOSU] UIEW JO UO JS] pue YUH Bulsojoue 


-ISod jsesojg jo Buiddeusno eueiquen 


(репициоэ) 


(senunuos) 


(a-voL 


“HL `$614) Apnis sy] едичэл [EUSA juasqe Juosaid с odA] (pasnj) ‚  erebinejo siey] sepIounyy 
(GL 'B14) Apnis sy | ¡e1Juan [EUSA juasqe ]Juasa.d с edA] (рэзп!) | 2n1s9/ зпиез$ем эерне$$ем 
(vz ‘By :1661) 
eÁexsuIpalA 9 JOJUB y ¿ g ¿ (2 de с IPJeyaıng easy  эерщедитоЭ 
(7, `614) Apnis siy [EUSA езиэл juasqe ]uasa.d с edA] (pasny) | exdiuos esıydnZ3 = =epiequinjoo 
(£L :B14) Apnis siy Je JUS A едиэл juesge Juosaid с odA] (pasnj) | eee euynedef эерциээпа 
epodonseßoaN 
esıuodel 
(2002) ‘1e Je meses Е = - - - 0 ejilydolajsy эершиий= 
(6-8 
‘7 "SB :G961) Weyelo ыедиэл едиэл juesge ¿ ¿ с eulyjuel eulyjuer Seplulyjuer 
ri esso|fus]d 
< (а зтезиди! 
T —9)z| ‘s6i4) Apnis SIUL  JeJejej sano [EUSA Juosald JU3saJd с od} С SIQIojNdlas эерцацшлэдл 
a (a 
oS —Yz | '$61-) Apnis $41 juasqe ¡eslop juasaJd juasaJd с edÁ] с гэшоэ xıuoddıy Seploiuoddiy 
sısuguoba1o 
=) (LL 614) Apnis sy] черэшлэци елчэл Juosald jJuasaJd с dA} Z UOJLISN{ эершечеч 
2 (a snuenyn| 
© —901 ‘s614) Apnıs siy | ¿juesqe ¡esop Jussald juasaJd с adÁl Z SngqwonS SEPIIWOAS 
(y sisuedny 
‘By ‘ec ‘Id :2961) sineg é é é é é Z  вивюшоцоио SepIqoJpÁH 
(s € euepide] 
'sB ‘ZL ‘Id :1961) sINeg ¿ ¿ & & E E sısdoyewog eeplIqoIpÂH 
(g eojuodel 
—01 ‘SBi4) Apnis $141 E [EUSA juesge jueseld e edA] С вэиил$$5у  Sepiaulissy 
ELUAPIP 
(a-96 '$614) Apnis siu | juasqe ¡esop juasaJd juasaud с adÁ] г хе/пе$$0]=) эерэцем 
SIUF19J94 Sspsnw $19]$|04 $19]$104 JE] J9]S|0Q ÁBOJOISIH $191$10а Je¡npe.l saridads exe | JOUBIH 
JoyewixoJdde Jepnpel -пре/ Jo sabe] Jeinpe4 JO зэбещцаеэ 
езиэл Jo зэбецаеэ - eo меш JO e6e]189 jo JOQUINN 
jo UOIHISSU]  шешю UON ye; pue зубы Bulsojoue 
-Isod ]sasojy Jo BuiddeyienQ  euelquen 
en 
LO 


(репициоэ) 


29 


RADULA-SUPPORTING STRUCTURES IN GASTROPODA 


(senunyuoo) 

(99861) seunidzsey 
(9861) эриоз 
(29861) Jeunidzse} 
(20661) 18puoq 
Apnjs $14. 

(2 By 
:0661) JOIN $ JolÄeL 

(vz ‘| ‘sby 
:1661) 1011 9 JOJUeY 


(VEL “BY :1661) sojuey 
(os ‘Bu :1661) sojUey 
(с ‘61 :Z661) 
чэАмэзелен $ JOJUEY 
(V2 BY :z/6L) лэриоа 


(al ‘бу :Z261) 18puoq 
(4-391 

11 "sB14) Apnis $141 

(IZ “Bu :0/6L) Japuod 


(az ‘бу :0461) лэрчоа 


99U9.19/19Y 


¿ 2 bs 

2 e E 

(3 is & 

E ¿ & 

¿ é ¿ 

¿ E с 

à ¿ & 

¿ 2 & 

E ¿ ¿ 

¿ e ¿ 

¿ едчэл éjuesqe 

езиэл елчэл juesge 

é é E 

é é © 
spsnw sJ91S|0Q sJ91S|0Q JE] 

Joyewixoidde Jejnpe. -прел Jo sabe] 
[eJJUSA Jo зэбецаеэ -11e9 иеш 
jo UOILISSU] UIEU JO чо} yo} pue зчбы 


-1504 159500) JO быааенэло 


©. ©: 


С. 


élueseld 


Juosa1d 


J31S|0Q 
лепрел 
JO 988/1989 
Buisojous 
3UBIQUIN 


2 0 euoeuw epyuido 
Маречц 

& 0 SIQIOPIDE/E) 
smebaeA 

E 0 

edoseyoo 

¿ 0 B/NISIPOJIIPY 

- О  sneelqe snuo) 

¿ (рэзп;) |  sn/noeq EJMSEH 
snIxed 

с edA} С PAASHIS 

& с ejjIoue елуюрпа$ 

é с 5$129404 EJJSAIIO 
eueyseje 

¿ € едиио] пол 

& 2 Маррер 107e/n984 
вполэатеа 

é С в//э}ебн]$ 

с edA] с elgez eyjstebuls 
eulja3snul 

¿ (pasny) | EULIEAJOA 
eaewbAd 

é (pasny) | ejsuıboseyy 

ABojoJsıH sJo]sjoq лепрел sardads 
JO зэбееэ 
JO JOQUINN 


SEPIPIIJJEMN 


эерчлорюею 


SNIEI[BH эерюиоэицолу 


эер! 59910 
ецэче.ао.ээн 
эершоЭ 
aepiqala] 
эер!а$194}$ 
эермлнорпэ$а 
эерШелно 


2RPIJJILUONIOA 
эериушоуюл 


Sep 
SEPHIIM 
eepl|euIBlen 


эершэешблеи/ 


exe | JOUBIH 


(penunuos) 


KATSUNO & SASAKI 


56 


(zz euelpjogals 
Y | '$6!-) Арп]$ siy ¡e1Juan [EUSA juasqe jJuasaJd 9 adÁ] (pasni) | ezoadsep ejsnoy sepiueeqApeig 
штепэпиал 
(LZ ‘FL ‘s6i4) Apnis sy] EUSA елзиэл juesqe juasaJd 9 adÁ] Z "JO “ds BIuoJad4 эерирщэчо 
syeubejs 
(oz 614) Apnis sy] ¡e1Juan [EUSA juasqe juasa.d 9 adÁ] (pasny) | eoeuuwAÁ7 aepieeuwA7 
(vz By 
:9007) Jaula3suayny ¡e1Juan [EUSA juesge juasaJd 9 adÁ1 с NUOSSNÓ еше  2epiieuoudis 
esıuodel 
(61 614) Apnis sy | [EUSA [EUSA juasqe Juosald 9 adh} (pasny) | eneuoydis  sepiieuoydis 
(3 ела} 
-38| ‘s6i4) Apnis $41 juesge [EUSA juesqe juasaJd 9 adÁ1 С $иоро/эз АН эерриорошоц9Э 
(а 
—981 `$614) Apnis SIUL juesqe - juasqe juasqe 9 adÁ] 0 elajnoo elsÁidy эерн$А!А\ 
(а ejyeuioun 
—V793| `$61-1) Apnis sy] juasqe - juasqe juasqe 9 adh} 0 BIUIJOACD эернинолеЭ 
(4 eoijuodefl 19 
—9/1 ‘s614) Apnis sy juesge - juasqe juasqe 9 edA] 0 pueydessouoddiny aepluyaio 
(a 
921 "sBI4) Ápn]s $141 JUSSQe JUSSQe JUSSQe 9 edA} 0 eejueble эиича эери!!Ча 
(а 
— ¿| '$6!-) Apnis $141 Jussqe - Jussqe juesge 9 odA] O  eoiuodel eojeyH эерэоицшен 
| epian]ad 
(30661) Jepuod e & & & & 0 BAJSOMUIOF  SEPHISONUIOT 
eEJEULICDOL] 
(2861) Jopuod é é é é é 0 eulyjeuy  SeplUIy}euy 
ээцэлэ!эч 2[9SNU SJ9]S|oq SI9]S|0Q JE] Ja\sjoq ÁBOJOISIH SI9]S|0q Je¡npe, sendads exe | JoyußIH 
Jojeuixoidde Jejnpes -пре/ Jo sabe] лепрел JO зэбеиеэ 
¡e11uen JO S9Be]nieo -1 eo меш JO 968/19 jo JOQUINN 
jo uoniesuj  UIELU JO uoN Je] pue зубы Buisojous 
-ISod jsasojg jo Buiddeusao  eueiquen 
(penunuos) 


MALACOLOGIA, 2008, 50(1-2): 57-173 


BIVALVIA OF THE DEEP ATLANTIC 


John A. Allen 


University Marine Biological Station, Millport, Scotland, KA28 OEG, 
and Woods Hole Oceanographic Institution, Massachusetts, U.S.A. 02543; 
jallen@udcf.gla.ed.uk 


ABSTRACT 


This paper lists the species and the distribution of the bivalves collected from the deep- 
sea expeditions undertaken by American, British and French research vessels in the At- 
lantic over a period of twelve years. Samples were taken from eleven basins, and the 
analysis is resticted to samples taken with the epibenthic sled from depths ranging from 
500 m to 5,000+ m. A preliminary analysis is made of the changing distribution with depth 
and discussion as to why some genera and families are either restricted to or are more 
dominant in the deep sea as compared with those found at shelf-sea depths. It provides a 
baseline of information against which future deep-sea sampling can be compared. 


Key words: deep sea, Bivalvia, Atlantic. 


INTRODUCTION 


During the years 1962 and 1974, various 
expeditions made by American, British and 
French scientists sampled the benthos of all 
but one of the major deep-sea basins of the 
Atlantic Ocean (Fig. 1). The exception was the 
Norwegian Basin, which was later investigated 
by Bouchet & Warén (1979). These expedi- 
tions sampled at depths ranging from the shelf/ 
slope break to the the greatest abyssal depth. 
They used a variety of sampling gear and hav- 
ing sorted the contents into the various phyla 
they distributed these to various experts. 

In the case of the bivalves, these were ex- 
amined for the most part by the late H. L. Sand- 
ers who led the American expeditions and by 
J. A. Allen. After sorting to species, work first 
concentrated on the distribution and functional 
morphology of species of the Protobranchia, 
which are dominant in the deeper samples. 
This work was recorded in the series of pa- 
pers cited below and culminated in an account 
of the zoogeography, diversity and origin of 
the deep-sea protobranch species of the At- 
lantic (Allen & Sanders, 1996b). Since then, 
attention has focused on the lamellibranch 
bivalves and, although there are a number of 
species that remain to be described (includ- 
ing protobranchs), an account of the Bivalvia 
as a whole can now be made. 

As in the earlier paper (Allen & Sanders, 
1996b), analysis is restricted to those samples 


D 


taken by the epibenthic sledge (Hessler & 
Sanders, 1967). This is because these 
samples, by far, provide a sufficient number 
of specimens/haul to make meaningful con- 
clusions as to dominance and distribution of 
species ocean wide. 

Information as to the species contained in the 
epibenthic sledge samples are listed in the Ap- 
pendices 1-3 and in descriptive accounts of in- 
dividual species in various papers stemming 
from the total collections (Allen, 1998, 2000a, 
b, 2001, 2004, in press; Allen & Hannah, 1989; 
Allen & Morgan, 1981; Allen & Sanders, 1966, 
1969, 1973, 1982, 1996a & b; Allen et al., 1995; 
Allen & Turner, 1974; Oliver & Allen, 1980a, b; 
Payne &Allen, 1991; Rhind & Allen, 1992; Sand- 
ers & Allen, 1973, 1977, 1985; Schein, 1989). 


COMPOSITION OF THE DEEP-SEA 
BIVALVE FAUNA OF THE ATLANTIC 


The distinction of bivalve families and sub- 
families changes with time and authority, but 
the most recent count indicates that of 108 liv- 
ing marine bivalve families some 87 families 
are present in the Atlantic. On the present data, 
between the shelf edge (200 m) and 500 m 79 
families are recorded. In the next 500 m there 
is a sharp drop in the number to 37 families 
present and thereafter a slow diminution in 
number with 28 recorded at 3,500 m and then 
a sharper drop to 14 at 5,000 m (Fig. 2). 


08 ALLEN 


FIG. 1. The deep-sea basins of the Atlantic. ANG — Angola; 
ARG —Argentina; BRA - Brazil; CAN — Canaries; CAP — Cape; 
CAV — Cape Verde; GUI — Guinea; NAM — North America; NFD 
— Newfoundland; NOR — Norwegian; SLE — Sierra Leone; SUR 
— Surinam; WEU — West European. 


When the species composition of the fami- 
lies in the deep-sea (500-5000 m) (not includ- 
ing the insertae cedis) is analysed, of the 456 
species and subspecies recorded, 133 (29%) 
are protobranchs and 323 (71%) are lamelli- 
branchs (Appendix 1). The percentage of proto- 
branch species increases with increasing depth 
from an average of 21.7% at 500 m to 57.3% 
at 4500+ m. These figures contrast with inter- 
tidal and shelf species, which in the case of 
Britain, 15 (6%) are protobranchs and 236 
(94%) are lamellibranchs. 

The lamellibranchs of the deep Atlantic are 
restricted to relatively few families with four — 


Pectinidae, Thyasiridae, Cuspidariidae, Verti- 
cordiidae — representing the overwhelming 
majority of species (Table 1). Many of the other 
families are restricted in the sense that only one 
or two genera of an otherwise common shal- 
low-water lamellibranch family may be present 
in the deep sea, for example, Dacrydium 
(Mytilidae) and Abra (Scrobicularidae). Similarly, 
of the protobranch families, two (Nuculanidae, 
Yoldiidae) dominate in terms of number of spe- 
cies, with species of two genera (Ledella, 
Yoldiella) dominating (Table 1). 

The geological records of these families are 
predominantly early, with ten from the Palaeo- 


BIVALVIA OF THE DEEP ATLANTIC 99 


80 


60 


40 


Number of families 


20 


500 1500 2500 3500 4500 
Depth (m) 


FIG. 2. Numbers of bivalve families found in the Atlantic at different 
depths. This is based on the present studies. 


TABLE 1.The bivalve families represented at depths between 500-5,000 m in the Atlantic, the num- 
ber of species recorded for each family and the earliest geological record of the family as given in 
Newell (1969). 


No. of Earliest No. of Earliest 
Protobranchia Species Geological Record Lamellibranchia Species Geological Record 
Solemyidae ‘i Ordovician Arcidae 5 Triassic 
Nucinellidae 1 Permian Limopsidae 10 Triassic 
Pristiglomidae 7 Recent Anomiidae 1 Cretaceous 
Nuculidae 3 Ordovician Mytilidae 13 Devonian 
Tindaridae =) Pliocene Pectinidae 38 Triassic 
Neilonellidae 9 Cretaceous Propeamussidae г. Permian 
Lametilidae 3 Recent Limidae 12 Carboniferous 
Nuculanidae 28 Devonian Ostreidae 1 Triassic 
Yoldiidae ar Cretaceous Lucinidae 3 Silurian 
Phaseolidae 4 Recent Thyasiridae 79 Triassic 
Siliculidae 3 Pliocene Montacutidae 12 Eocene 
Malletiidae 10 Ordovician Neoleptonidae 1 Pliocene 
Carditidae 3 Devonian 
Cardiidae 4 Triassic 
Mactridae 1 Cretaceous 
Scrobicularidae 8 Cretaceous 
Astartidae 6 Devonian 
Kelliellidae 12 Eocene 
Vesicomyidae ? Miocene 
Veneridae 2 Cretaceous 
Hiatellidae 2 Jurassic 
Xylophagidae 1 Cretaceous 
Teredinidae 1 Paleocene 
Pholadomyidae 2 Triassic 
Lyonsiidae 1 Eocene 
Thraciidae 10 Jurassic 
Periplomatidae 1 Cretaceous 
Poromyidae 6 Cretaceous 
Cuspidariidae 54 Cretaceous 
Verticordiidae 27 Paleocene 


60 ALLEN 


TABLE 2. The number of species, living specimens, and endemism of bivalves taken between 1962 
and 1974 using the epibenthic sled from the deep-sea basins of the Atlantic. 


No. of 
Basin Stations No. of Spp. 
North America 49 52,491 
Surinam 15 16815 
Brazil 7 1,452 
Argentine 16 14,856 
Cape Verde 3 299 
West European 212 81,493 
Canaries 9 699 
Sierra Leone 11 2,433 
Guinea 12 379 
Angola 21 107141 
Cape 27 28,832 


zoic, 20 from the Mesozoic, and 12 from the 
Cenozoic. Ofthese the geological history ofthe 
protobranchs is particularly interesting in that 
of the 12 families five are from the Mesozoic 
(three Recent), two from the Cenozoic, and five 
are from the Paleozoic (3 Ordovician) reflect- 
ing both the antiquity of the subclass and the 
recent evolutionary expansion of species in the 
deep-sea (Allen & Sanders, 1996b) (Table 1). 

In general, the greater the number of 
samples taken in a particular basin the more 
species are recorded, with the number of rarer 
species increasing. Thus, in the case of the 
West European Basin, the most sampled ba- 
sin with 112 epibenthic trawl samples taken, 
some 203 species were recorded, while in the 
case of the North America Basin the next most 
sampled (49), 193 species were recorded. The 


NO & > 
© о © 


No. species/sample 


BEN 
(<>) 


10 100 
No. specimens/sample 


No. of Spp. & No. of Endemic 
Subspp. Spp. & Subspp. % Endemic 

193+4 57 28.9 
129+3 32 24.2 
49 8 16.3 
116+3 39+1 336 
9+1 1 al 
203+8 70+3 34.6 
49+2 6 li 
67+3 2 2.9 
37+1 2 5.9 
123+4 35 27.6 

43+3 2 


plots from the North America and West Euro- 
pean basins would indicate that, subject to the 
sampling efficiency of the epibenthic sled, at 
least 15 samples are required at any particu- 
lar 500 m depth interval to record the number 
of species present with any degree of accu- 
racy. Taking into account the sampling inten- 
sity, analysis suggests that approximately 30% 
of the species present are endemic to any 
basin (Table 2, Appendix 1). 

As would be expected, when the number of 
species at each station is plotted against the 
total number of specimens present, the num- 
ber of species increases with increasing num- 
bers of specimens (Fig. 3). However, the 
maximum number of species present peaks 
when more than 100 specimens are present 
in the sample (Fig. 3). In samples containing 


1000 


FIG. 3. The number of species present in each epibenthic sledge sample 
taken from the Atlantic basins compared with the number of specimens in 


each sample. 


BIVALVIA OF THE DEEP ATLANTIC 61 


2864m N=716 

60 

50 
2 40 a 
Е E 
9 481m N= 1446 E 
© 0 
(0) Ф 
O O 
о 9 
E E 
(0) DO 
SER O 
© © 

20 

10 


CAPE 


5 10 15 20 20 
Individual species 


60 


50 


4980m N = 1411 


40 
| 923m N=2701 


30 


20 


\ SURINAM 
SAS 


Een — 


Individual species 


FIG. 4. Examples of plots of individual species against descending percentage occurrence in indi- 
vidual samples at upper slope and abyssal depths in different deep-sea basins. 


100+ specimens although the minimum the 
number of species present increases from < 
10, the maximum number is rarely more than 
35. In only one sample containing more than 
9,000 specimens were 43 species recorded 
(Fig. 3). This pattern of occurrence is true for 
all the basins sampled. 

Analysis of individual samples with more than 
100 specimens shows that there is a pattern 
in which five species dominate each sample 
with the remainder having rapidly diminishing 
percentage representation. This pattern which 
is seen in all samples in all basins of the Atlan- 


tic at all depths (Figs. 4, 5). Typical examples 
are shown in Fig. 4, in which upper slope and 
abyssal samples are compared in basins from 
opposite sides of the mid-Atlantic ridge and 
from north and south of the equator. Taking all 
basins into consideration the extreme limits of 
percentage occurrence of the first dominant 
species lies between 11.8% and 95.7% of a 
sample. Taking all samples containing > 100 
specimens, the average percentage occur- 
rences of first five dominants in descending 
dominance are 44.9%, 16.1%, 10.6%, 7.1% 
and 3.7% respectively, that is, representing an 


62 ALLEN 


No. of samples 
if «2 OF. 40-23 


100 35 18 4 13 


80 


60 


40 


Percentage spread 


20 


(BS) M 
10.6 
7.1 
SM. 


NAM SUR BRA ARG WEU WEU SLE ANG CAP 


ab f 


FIG. 5. The percentage spread of the first five dominant species in samples containing 
100+ specimens in individual Atlantic basins at 500-meter intervals in depths ranging 
from 500 to 5,000 meters. NAM - North America, SUR — Surinam, BRA - Brazil, ARG — 
Argentina, WEUab — West European (American and British samples), WEUf — West 
European (French samples), SLE — Sierra Leone, ANG — Angola, CAP — Cape. 


average of 76.4% of the specimens present in 
a sample. It is also typical that the percentage 
decline in the second to fourth dominants is 
less steep than that between the first and sec- 
ond dominants and the fourth and fifth (Fig. 4). 
The averages of the first five dominants is also 
remarkably constant for all the basins of the 
Atlantic (Fig. 5), nor does the degree of domi- 
nance relate to depth. High percentage domi- 
nance can be observed in samples at 500 m 
as well as at 4,500 m (Fig. 4). What does vary 
with depth are the particular species that com- 
prise the dominants. 

Of the 456 species listed only 57 are first 
dominants and 122 are first, second or third 
dominants (Appendices 1, 2) at some depth 
within their individual depth range. Of the first 
dominants, 33 are protobranchs and 24 are 
lamellibranchs, and of the first three dominants 
62 are protobranchs and 60 lamellibranchs of 
which latter one third are thyasirids. Of the 
common families in the deep sea, the Cus- 
pidariidae are least represented among the 
dominant species despite their ubiquity. Indi- 
vidual depth ranges of the dominants varies 
but, summing all basins (i.e., a species may 
be a dominant in more than one basin and 
may have a different depth range in each), 
72% dominants have depth ranges of less than 


1,500 m and 82% have ranges of less than 
2,000 m. (Note, these percentages include 
15% that occur at one station only.) Less than 
9% of the dominants have depth ranges of 
3,000+ m, and these comprise ten species, of 
which four are protobranchs, four are thyasi- 
rids and one kelliellid and one cuspidariid. 

Of the 11 basins sampled, no one species was 
recorded in all (Table 3). However, in the case 


TABLE 3. The number of species and subspe- 
cies present in one and present in two or more 
of eleven Atlantic basins. 


No. of Basins No. of Species 


2000 YXO00+20Nn—- 
— 
N 


— — 


BIVALVIA OF THE DEEP ATLANTIC 


No. of samples 
eas 40. 25 12 16 21 35 


20 


15 Г 


Av. no. species/sample 


ас 


500 2500 4500 
Depth (т) 


FIG. 6. The average number of species п samples with 
100+ specimens taken from all the deep-sea basins of 
the Atlantic in 500-meter intervals between depths of 
500 to 4,500 meters (T). Below these are further di- 
vided into the various feeding types. С: commensal; 


63 


D: deposit; F: filter; M: macrofeeders. 


of some of the basins, particularly the Cape 
Verde and Brazil, few samples were taken. It is 
seems likely that widely distributed species re- 
corded in a least seven basins, that is, the 
greater part of the deep Atlantic, occur through- 
out the ocean. Taking this as the base line, 28 
species were recorded in seven or more ba- 
sins, and of these all but six dominated in one 
or more basins. The most common dominant, 
Ledella ultima, was recorded in 10 of the 11 
basins sampled and was recorded as a first 
dominant in five basins and second dominant 
in a further two basins at abyssal depths rang- 
ing from 3,828 m to 5,280 т (Appendices 1-2). 

Of 65 species that occur in just two basins, 
52% occur in north/south adjacent basins on 
one or other side of the mid-Atlantic ridge. In 
contrast, only 23% occur in east/west adjacent 
basins that are separated by the mid-Altlantic 
ridge, for example, North America-West Euro- 
pean and Argentine-Angola/Cape. The remain- 
ing 25% occur distant from each other but show 
no predominance to one side of the mid-Atlan- 
tic ridge or the other and not one is a dominant 
species. This latter probably relates to rarity and 
to degree of sampling effort. 

Analysis of dominance at 500-m intervals 
between 500 m and 4,000+ m. confirms the 
general increase in the percentage of 


protobranch species with increasing depth and 
the much higher percentage of dominant 
protobranch species as compared with lamel- 
libranchs at all depth intervals. The increase 
in the number and species of protobranchs 
with increasing depth is a reflection of the in- 
crease in deposit feeding species with depth 
and, of which, protobranchs are by far the 
major proportion (Fig. 6). 

The species can be divided into four main 
feeding catagories, namely deposit, filter, 
macrofeeders, and commensals (which may 
be filter or macrofeeders) (Fig. 6). The aver- 
age number of species in each catagory for 
samples containing 100+ specimens within 
500 m depth intervals between 500 m to 4,500 
m shows that suspension feeders decrease 
with increasing depths below 1,000 m. In con- 
trast, there is an increase in deposit feeders 
down to a depth of 2,500 m and thereafter a 
levelling off of species numbers. Macrofeeders 
are a small but persistant group at all depths 
with a slight diminution in numbers as depth 
increases. Commensal species are few in 
number but do occur at all depths. It is likely 
that these latter are underestimated simply 
because many remain with their non-mollus- 
can hosts and await discovery by experts deal- 
ing with the host species. 


64 ALLEN 


DISCUSSION 


Species richness and biodiversity in the deep 
sea have been the subjects of a number of 
papers in recent years, for example, Allen & 
Sanders (1996b), Levin et al. (2001), Snelgrove 
& Smith (2002), Stuart et al. (2003), and Rex 
et al. (2005), and information to support these 
works has stemmed in part from the bivalve 
data presented here. This said, it is not the 
purpose of the present paper to further discuss 
diversity, more it is the presentation of basic 
information from wide ranging international 
sampling programmes of an ocean that are 
unlikely to be repeated in the forseeable fu- 
ture. The majority of the bivalve samples are 
stored in the Muséum National d’Histoire 
Naturelle, Paris, and the Museum of Compara- 
tive Zoology, Harvard University, Cambridge. 

The main environmental factors that control 
the occurrence and evolution of the bivalves 
in the deep-sea are food supply and the effect 
of high pressure on the physiology of body 
functions. Although it is now known that the 
sediments are diversified and that there is 
seasonal organic input from surface waters, 
particularly in higher latitudes, which is influ- 
enced by internal waves and down-slope sedi- 
ment slumping (Levin et al., 2001; Stuart et 
al., 2003; Rex et al., 2005), the available food 
supply differs markedly in its composition to 
that in shallow waters. Particles arriving at the 
sea bed have high scleroprotein content and 
are much reduced in quantity (McCave, 1974). 
This impoverished food supply is probably the 
reason for small body size, with most deep- 
sea bivalves being < 1 cm in length and the 
majority < 5 mm in length. It also explains why 
there is a large increase in the number of de- 
posit feeding species. In addition, it should be 
noted that many deposit feeding protobranch 
species at abyssal depths have increased gut 
length compared with their shallower water 
relatives (Allen, 1992) and also, the few 
tellinacean species present have modified in- 
testines of increased length and volume (Allen 
& Sanders, 1966). Both allow increased time 
for digestion. 

Associated with small size is a reduction in 
the size of the gills. This may involve partial or 
total loss of the outer demibranch which may 
also be partly explained by the effect of high 
pressure in relation to respiratory require- 
ments. Nevertheless, the gross internal mor- 
phology of the deep-sea species does not 
differ markedly from their shallow-water coun- 
terparts. 


Whereas the ecology and evolution of the 
protobranchs of the deep Atlantic have been 
discussed at some length (Allen & Sanders, 
1996b), that of the deep-water lamellibranchs 
is less well known. Descriptions of many lamel- 
libranch species have been detailed (Allen, 
1998, 2000a, b, 2001, 2004; Allen & Morgan, 
1981; Allen & Turner, 1974; Oliver & Allen, 
1980a, b; Payne &Allen, 1991), but their ecol- 
ogy and adaptations to abyssal life are much 
less well-known. It has been pointed out that 
the number lamellibranch families in the deep- 
sea is small and restricted to a very few spe- 
cies and some to a singl 

e genus. Some groups, such as the 
septibranchs, have highly specialized macro- 
feeding habits and others may have 
specialised commensal relationships or hab- 
its, such as that of Adipicola and /dasola, which 
are restricted to the remains of whales. It is 
interesting that both of these groups have more 
or less the same percentage occurrence in the 
deep-sea as they do in shallow-water (Fig. 6). 
Nor do the deep-sea representatives of the 
two groups have any great anatomical differ- 
ences from their shallow-water counterparts. 
Presumably, their food sources are much the 
same at whatever depth of their occurrence. 

A similar argument might be made of the 
pectinaceans, in particular the Propeamussi- 
dae, mobile carnivors that are well-represented 
in the deep-sea. These show some structural 
modification in that not only are they small in 
size, but their shells are extremely thin and 
delicate and adapted for a life at the surface of 
the nephaloid layer. It would appear likely that 
deep-sea species spend much of their time 
swimming just above the sediment surface and, 
being mobile, activily seek out their food. Also 
of significance is that the Propeamussidae may 
be living relicts of a group thought to have be- 
come extinct at the end of the Paleozoic pe- 
riod (Waller, 1970, 2006). 

The most dominant lamellibranch family 
present in the deep-sea is the Thyasiridae. 
Other lucinoideans excepted, and of which the 
lucinids and ungulinids are largely restricted 
to depths of < 1,000 m, the thyasirids have a 
morphology that is unlike that of other bivalves 
(Allen, 1958; Payne & Allen, 1991). Charac- 
teristically they have foot that is modified and 
used to form an inhalent tube to the surface 
for the intake of food and water. In addition, 
they have an unusual visceral mass in which 
gonads and digestive diverticula are largely 
restricted to lateral lobes on either side of the 
body. It is difficult to to think why this latter 


BIVALVIA OF THE DEEP ATLANTIC 65 


should be advantageous to life at great depths, 
but the unusual structure and function of the 
foot may well be. Unlike the shallow water 
lucinids and thyasirids which have thickened 
gills containing symbionts (Southward, 1986; 
Taylor & Glover, 2000), most of the deep-sea 
thyasirids are without (Payne & Allen, 1991). 
Presumably this is because the majority does 
not live in environments deficient in oxygen. 

The deep-sea representatives of suspension- 
feeding lamellibranch families that are found 
at all depths (Kelliellidae, Arcidae, Limopsidae 
and Limatulidae) differ little from their shallow- 
water representatives. Other than small size 
and reduced gill size, there appear to be no 
modifications for life at great depths. In fact 
some, such as species of Kelliella, are first 
dominants and occur in very large numbers. 
This is also true of Dacrydium, one of very few 
mytilids to occur in the deep sea yet, apart from 
small size, it shows no anatomical differences 
that could account for their success. It is likely 
that explanation relates to physiological rather 
than gross anatomical differences. 

Of all the lamellibranch families in the deep 
sea the Thyasiridae is the one that shows the 
most remarkable degree of speciation and a 
great many species remain to be described 
(see Appendix 2). Whether this is because that 
there is a paucity of competition in deep wa- 
ters from the more recently evolved and more 
successful families that dominate in shallow 
waters, for example, veneraceans and 
tellinaceans, is debateable. However, it would 
appear that these more recently evolved and 
successful families in shallow water have dis- 
placed down-slope species of more ancient 
origin (Allen, 1996). It has also been recently 
suggested that molluscs have penetrated the 
deep sea by a source-sink mechanism (Rex 
et al., 2005). Nevertheless, there are 
undoubedly resident deep-sea species that are 
restricted to the abyss (e.g., Ledella ultima), 
many with oceanwide distributions and which 
successfully reproduce at those depths. Most 
of the species restricted to lower bathyal and 
abyssal depths are protobranchs but a few 
lamellibranch species are restricted to these 
depths and include Limopsis tenella, 
Dacrydium abyssorum, and Thyasira inflata. 
Furthermore, judging by their restricted depth 
distributions this also applies to bathyal spe- 
cies at mid- and lower-slope depths. This is 
not to say that the sink-source mechanism 
does not apply to species with a wide distribu- 
tion extending from shallow to deep water (Ap- 
pendix 2). 


ACKNOWLEDGEMENTS 


| would like to thank Dr. Thomas R. Waller of 
the Department of Paleobiology, Smithsonian 
Institution, Washingtion, D.C., for his identifi- 
cations of the pectinaceans, without which this 
study would have been seriously incomplete: 
also, to Dr. C. Miller of Oregon State Univer- 
sity whose critical analysis of the original draft 
was so invaluable. There are also the detailed 
comments and suggestions from the referees 
which were much appreciated. | must give trib- 
ute to all the hard work of my colleagues and 
the ships crews that made the deep-sea ex- 
peditions so successful. Finally many thanks 
must go to Miss. Tracy Price for her hard work 
in ensuring that the manuscript met all the 
detailed editorial requirements. 


LITERATURE CITED 


ALLEN, J. А., 1958, On the basic form and ad- 
aptations to habitat in the Lucinacea 
(Eulamellibranchia). Philosophical Transac- 
tions of the Royal Society of London, 241: 421-— 
484. 

ALLEN, J. A., 1992, The evolution of the indgut 
of the deep-sea protobranch bivalves. Ameri- 
can Malacological Bulletin, 9: 187-192. 

ALLEN, J. A., 1998, The deep-water species of 
Dacrydium Torrell, 1859 (Dacryniinae: 
Mytilidae: Bivalvia) of the Atlantic. Malacologia, 
40: 1-36. 

ALLEN, J.A., 2000a, An unusual montacutid from 
the deep Atlantic. Journal of the Marine Bio- 
logical Association of the United Kingdom, 80: 
827-834. 

ALLEN, J. A., 2000b, A new deep-sea species 
of the genus Neolepton (Bivalvia: Cyamoidea: 
Neoleptonidae) from the Argentine Basin. 
Malacologia, 42: 123-129. 

ALLEN, J. A., 2001, The family Kelliellidae 
(Bivalvia: Heterodonta) from the deep Atlantic 
and its relationship with the family 
Vesicomyidae. Zoological Journal of the Lin- 
nean Society, 131: 199-226. 

ALLEN, J. А., 2004, The Recent species of the 
genus Limatula and Limea (Bivalvia, Limacea) 
present in the Atlantic, with particular reference 
to those in deep water. Journal of Natural His- 
tory, 38: 2591-2653. 

ALLEN, J. A. & F. HANNAH, 1986, A reclassifi- 
cation of the Recent genera of the subclass 
Protobranchia (Mollusca: Bivalvia). Journal of 
Conchology, 32: 225-249. 

ALLEN, J. A. & F. HANNAH, 1989, Studies on 
the deep-sea Protobranchia. The subfamily 
Ledellinae (Nuculanidae). Bulletin of the Brit- 
ish Museum (Natural History), Zoology, 55: 
123-171. 

ALLEN, J. A. & К. E. MORGAN, 1981, The func- 
tional morphology of the families Cuspidariidae 


66 ALLEN 


and Poromyidae (Mollusca: Bivalvia) of the 
abyssal Atlantic. Philosophical Transactions of 
the Royal Society of London, В, 291: 413-546. 

ALLEN, J. A. 8 H. Е. SANDERS, 1966, Adapta- 
tions to abyssal life as shown by the bivalve 
Abra profundorum (Smith). Deep-sea Re- 
search, 13: 1175-1184. 

ALLEN, J. A. 8 Н. Е. SANDERS, 1969, Nucinella 
serrei (Protobranchia) a monomyarian 
solemyid and possible living actinodont. 
Malacologia, 7: 381-396. 

ALLEN, J. À. & H. L. SANDERS, 1973, Studies 
on the deep-sea Protobranchia. The families 
Siliculidae and Lametilidae. Bulletin of the 
Museum of Comparative Zoology Harvard 
University, 145: 263-310. 

ALLEN, J. À. & H. L. SANDERS, 1982, Studies 
on the deep-sea Protobranchia. The subfam- 
ily Spinulinae (family Nuculanidae). Bulletin of 
the Museum of Comparative Zoology Harvard 
University, 150: 1-30. 

ALLEN, J. A. & H. L. SANDERS, 1996a, Studies 
on the deep-sea Protobranchia (Bivalvia); the 
family Neilonellidae and the subfamily 
Nuculaninae. Bulletin of the British Museum 
(Natural History), Zoology, 62: 101-132. 

ALLEN, J. A. & Н. L. SANDERS, 1996b, The zoo- 
geography, diversity and origin of the deep-sea 
protobranch bivalves of the Atlantic: The epi- 
logue. Progress in Oceanography, 38: 95-153. 

ALLEN, J. A., H. Е. SANDERS & Е. HANNAH, 
1995, Studies on the deep-sea Protobranchia 
(Bivalvia); the subfamily Yoldiellinae. Bulletin 
of the British Museum (Natural History), Zool- 
ogy, 61: 11-90. 

ALLEN, J.A. & J. F. TURNER, 1974, On the func- 
tional morphology of the family Verticordiidae 
(Bivalvia) with descriptions of new species from 
the abyssal Atlantic. Philosophical Transactions 
of the Royal Society of London, B, 268: 401- 
536; 

BOUCHET, Р. Н. & А. WAREN, 1979, The mol- 
luscan fauna of the Norwegian Sea and its re- 
lation to other faunas. Sarsia, 64: 211-243. 

HESSLER, К. К. & H. L. SANDERS, 1967, Fau- 
nal diversity in the deep-sea. Deep-Sea Re- 
search, 14: 65-78. 

LEVIN, L. A., R. J. ЕТТЕК, M. A. REX, А. J. 
GOODAY, C. R. SMITH, J. PINADA, C. T. 
STUART, R. R. HESSLER & D. PAWSON, 
2001, Environmental influences on regional 
deep-sea species diversity. Annual Review of 
Ecology and Systematics, 32: 51-93. 

McCAVE, |. N., 1974, The benthic boundary 
layer. New York & London, Plenum Press. 

NEWELL, N. D., 1969, Classification of the 
Bivalvia. Pp. 205-224, in: R. С. MOORE, Trea- 
tise on invertibrate paleontology. Part N, 1, 
Mollusca 6, Bivalvia. 

OLIVER, G. & J. À. ALLEN, 1980a, The functional 
and adaptive morphology of the deep-sea spe- 
cies of the Arcacea (Mollusca: Bivalvia) from 
the Atlantic. Philosophical Transactions of the 
Royal Society of London, B, 291: 45-76. 

OLIVER, G. & J. A. ALLEN, 1980b, The func- 
tional and adaptive morphology of the family 
Limopsidae (Bivalvia: Arcoida) from the Atlan- 


tic. Philosophical Transactions of the Royal 
Society of London, B, 2901: 77-125. 

PAYNE, C. M. & J. A. ALLEN, 1991, The mor- 
phology of deep-sea Thyasiridae (Mollusca: 
Bivalvia) from the Atlantic Ocean. Philosophi- 
cal Transactions of the Royal Society of Lon- 
don, B, 334: 481-562. 

REX, M. А., С. В. McCLAIN, N. A. JOHNSON, 
R. J. ETTER, J. A. ALLEN, P. BOUCHET & A. 
WAREN, 2005, A source-sink hypothesis for 
abyssal diversity. The American Naturalist, 165: 
163-178. 

RHIND, P. М. & J. A. ALLEN, 1992, Studies on 
the deep-sea Protobranchia (Bilvalvia): the 
family Nuculidae, Bulletin of the British Mu- 
seum (Natural History), Zoology, 58: 61-93. 

SANDERS, Н. L. 8 J. A. ALLEN, 1973, Studies 
on the deep-sea Protobranchia (Bivalvia): pro- 
logue and the Pristiglomidae. Bulletin of the 
Museum of Comparative Zoology Harvard 
University, 145: 237-271. 

SANDERS, H. L. & J. A. ALLEN, 1977, Studies 
on the deep-sea Protobranchia (Bivalvia): the 
family Tindariidae and the genus Pseudo- 
tindaria. Bulletin of the Museum of Compara- 
tive Zoology Harvard University, 148: 23-59. 

SANDERS, H. L. & J. A. ALLEN, 1985, Studies 
on the deep-sea Protobranchia (Bivalvia): the 
family Malletiidae. Bulletin of the British Mu- 
seum (Natural History), Zoology, 49: 195-238. 

SCHEIN, E., 1989, Pectinidae (Mollusca, 
Bivalvia) bathyaux et abyssaux des cam- 
pagnes BIOGAS (Golfe de Gascogne) 
systématique et biogéographie. Annales de 
l’Institut Océanographique, n.s., 65: 59-125. 

SNELGROVE, Р. V. К. & С. В. SMITH, 2002, A 
riot of species in an environmental calm: the 
paradox of the species-rich deep-sea floor. 
Oceanography and Marine Biology: an Annual 
Review, 40: 311-342. 

SOUTHWARD, Е. C., 1986, Gill symbionts in 
thyasirids and other bivalve molluscs. Journal 
of the Marine Biological Association of the 
United Kingdom, 66: 889-914. 

STUART, С. Т., М.А. REX &R. J. ETTER, 2003, 
Large-scale spatial and temporal patterns of 
deep-sea benthic species diversity. Pp. 297- 
313, in: P. A. TYLER, ed., Ecosystems of the 
world: 28. Ecosystems of the deep oceans. 
Amsterdam, Elsevier. 

TAYLOR, J. D. & E. A. GLOVER, 2000, Func- 
tional anatomy, chemosynthesis and evolution 
of the Lucinidae. In: E. M. HARPER, J. D. TAYLOR 
& J.A. CRAME, eds., The evolutionary biology of 
othe Bivalvia. Geological Socieity of London 
Special Publicaion, 177: 207-225 

WALLER, T. R., 1970, The glass scallop 
Propeamussium, a living relict of the past. An- 
nual Report for 1970 of the American Malaco- 
logical Union, pp. 5-7. 

WALLER, T. R., 2006, Phylogeny of families fami- 
lies in the Pectinacea (Mollusca: Bivalvia): im- 
portance of the fossil record. Zoological Journal 
of the Linnean Society, 148: 313-342. 


Revised ms. accepted 8 October 2007 


BIVALVIA OF THE DEEP ATLANTIC 67 
APPENDIX 1 


Distribution of living deep-sea bivalves of the various basins of the Atlantic ARG, Argentine; 
BRA, Brazil; SUR, Surinam; CAV, Cape Verde: NAM, North America; WEU, West European; 
CAN, Canaries; SLE, Sierra Leone; GUI, Guinea; ANG, Angola; CAP, Cape. Presence of a 
species is indicated by + and when a sample or samples within a basin contain a species that 
ranks in the first three most common within the sample this is indicated by “d” (dominant). 


ARG BRA SUR CAV NAM WEU САМ SLE GU1 ANG CAP 


Nucinella pretiosa + + 
Gould, 1861 

Solemya grandis + 
Verrill & Bush, 1898 

Solemya sp. + 

Solemya sp. 195 # 

Solemya sp. 237 + 

Solemya sp. 259 + + 

Solemya sp. 280 + 

Solemya sp. 293 + 

Pristigloma alba # # + + # + + 
Sanders & Allen, 1973 

Pristigloma nitens + + + + d + + 
(Jeffreys, 1876) 

Pristigloma sp. a of 

Microgloma pusilla + 
(Jeffreys, 1879) 

Microgloma turnerae + d at 
Sanders & Allen, 1973 

Microgloma yongei d + + d Hr 
Sanders & Allen, 1973 

Microgloma sp. $ d + 

Nucula callicredemna + 
Dall, 1890 

Nuculidae sp. 280 + 

Nuculidae sp. 330 + 

Deminucula atacellana d + d d + + + 
(Schenk, 1939) 

Nuculoidea bushae + d d d + d + 
(Dollfus, 1898) 

Nuculoidea pernambucensis d 
(Smith, 1885) 

Nuculoma elongata + 
Rhind & Allen, 1992 

Nuculoma granulosa + d d 1 
(Verrill, 1884) 

Nuculoma perforata + d + 
Rhind & Allen, 1992 

Nuculoma similis + d 
Rhind & Allen, 1992 

Brevinucula subtrangularis + 
Rhind & Allen, 1992 

Brevinucula verrilli * + + + + d d d 
(Dall, 1886) 


(continues) 


68 ALLEN 
(continued) 


ARG BRA SUR CAV NAMWEU CAN SLE GU1 ANG CAP 


Tindaria miniscula + + d + + 
Sanders & Allen, 1977 

Tindaria callistiformis + + d + + + 
Verrill 8 Bush, 1897 

Tindaria hessleri + + + 
Sanders & Allen, 1977 

Tindaria perrieri Er 
(Dautzenberg & Fischer, 1896) 

Tindaria sp. 188 E 

Neilonella corpulenta + : 
(Dall, 1881) 

Neilonella guineensis + of + + + + + 
(Theile, 1931) 

Neilonella hampsoni + + 
Allen & Sanders, 1996 

Neilonella salicensis d d d + + d d + 
(Seguenza, 1877) 

Neilonella whoii + = d d + + + d 
Allen & Sanders, 1996 

Pseudotindaria championi + + 
(Clarke, 1961) 

Pseudotindaria erebus + + + + + + zb d 
(Clarke, 1959) 

Pseudotindaria sp. + 

Prelametila clarkei d + d 
Allen & Sanders, 1973 

Prelametila sp. 247 + + 

Lametila abyssorum d + si 
Allen & Sanders, 1973 

Phaseolus sp. a + 

Phaseolus sp. b + 

Phaseolus sp. c d 

Phaseolus sp. d + 

Tindariopsis aeolata + 
(Dall, 1890) 

Tindariopsis agatheda + + d 
(Dall, 1889) 

Tindariopsis sp. + 

Ledella aberrenta of + + + + 
Allen & Sanders, 1996 

Ledella acinula + d + + + 
(Dall, 1890) 

Ledella acuminata d + 
(Jeffreys, 1870) 

Ledella galatheae + 
Knudsen, 1970 

Ledella jamesi d d 
Allen & Hannah, 1989 

Ledella lusitanensis + 
Allen & Hannah, 1989 

Ledella oxira a; 
(Dall, 1927) 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 69 
(continued) 


ARG BRA SUR CAV NAMWEU CAN SLE GU1 ANG CAP 


Ledella parva + 
Verrill & Bush, 1897 

Ledella pustulosa argentinae d 
Allen & Hannah, 1989 

Ledella pustulosa hampsoni id $ 
Allen & Hannah, 1989 

Ledella pustulosa marshalli d + 
Allen & Hannah, 1989 

Ledella pustulosa pustulosa d 
(Jeffreys, 1876) 

Ledella sandersi + 
Allen & Hannah, 1989 

Ledella similis + 
Allen & Hannah, 1989 

Ledella sublevis d + ть + + d % 
Verrill & Bush, 1893 

Ledella ultima + + d d d d d + d d 
(Smith, 1885) 

Ledella sp. + 

Spinula filatovae + № d d + 
Knudsen, 1967 

Spinula hilleri Es + + E + + au + 
Allen & Sanders, 1982 

Spinula scheltemae + + d 
Allen & Sanders, 1982 

Spinula subexisa it * 
(Dautzenberg & Fischer, 1897) 

Spinula sp. + 

Nuculana acuta + + 
(Сопгаа, 1831) 

Nuculana commutata + 
(Philippi, 1844) 

Nuculana vestita + + d 
(Locard, 1898) 

Propeleda paucistriata + 
Allen & Sanders, 1996 

Propeleda carpenteri d 
(Dall, 1881) 

Propeleda louiseae + 
(Clarke, 1961) 

Yoldiella americana + d d 
Allen, Sanders & Hannah, 1995 

Yoldiella argentinensis + 
Allen, Sanders & Hannah, 1995 

Yoldiella artipica + + 
Allen, Sanders & Hannah, 1995 

Yoldiella biguttata + + + 
Allen, Sanders & Hannah, 1995 

Yoldiella bilanta + + d 
Allen, Sanders & Hannah, 1995 

Yoldiella blanda d 
Allen, Sanders & Hannah, 1995 


(continues) 


70 ALLEN 


(continued) 


ARG BRA SUR CAV NAMWEU CAN SLE GU1 ANG CAP 


Yoldiella biscayensis d 
Allen, Sanders & Hannah, 1995 

Yoldiella curta Е d d + d + + d + 
Verrill & Bush, 1898 

Yoldiella capensis ar d 
Allen, Sanders & Hannah, 1995 

Yoldiella dissimilis == + 
Verrill & Bush, 1898 | 

Yoldiella ella + ie d d d u; + 
Allen, Sanders € Hannah, 1995 

Yoldiella enata d + + 
Allen, Sanders & Hannah, 1995 

Yoldiella extensa d 
Allen, Sanders & Hannah, 1995 

Yoldiella fabula a + + + к 
Allen, Sanders & Hannah, 1995 

Yoldiella frigida + d d 
(Torrell, 1859) 

Yoldiella hanna d d 
Allen, Sanders & Hannah, 1995 

Yoldiella inconspicua africana + + + an 
Allen, Sanders & Hannah, 1995 

Yoldiella inconspicua d d 
inconspicua 
Verrill & Bush, 1898 

Yoldiella profundorum + 
Allen, Sanders & Hannah, 1995 

Yoldiella insculpta d + 
(Jeffreys, 1879) 

Yoldiella jeffreysi + + d d d + at 
(Hidalgo, 1879) 

Yoldiella lata d 
(Jeffreys, 1876) 

Yoldiella lucida EE d + 
(Loven, 1848) 

Yoldiella obesa obesa + + 
(Stimpson, 1851) 

Yoldiella obesa incala d 
Allen, Sanders & Hannah, 1995 

Yoldiella ovata d 
Allen, Sanders & Hannah, 1995 

Yoldiella perplexa d 
Allen, Sanders & Hannah, 1995 

Yoldiella pseudolata d + # 
Allen, Sanders & Hannah, 1995 

Yoldiella robusta + 
Allen, Sanders & Hannah, 1995 

Yoldiella similirus + a 
Allen, Sanders & Hannah, 1995 

Yoldiella similis + 
Allen, Sanders & Hannah, 1995 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC tA 
(continued) 


ARG BRA SUR CAV NAMWEU CAN SLE GU1 ANG CAP 


Yoldiella sinuosa + 
Allen, Sanders & Hannah, 1995 

Yoldiella subcircularis A d d + + + + 
(Odhner, 1960) 

Yoldiella valetta + sh + 
Allen, Sanders & Hannah, 1995 

Yoldiella sp. 246 at 

Yoldiella sp. 314 + 

Portlandia abyssorum + + it 
(Knudsen, 1970) 

Portlandia lenticula + + + 
(Moller, 1842) 

Portlandia minuta d ot 
Allen, Sanders & Hannah, 1995 

Serapta sp. + 

Silicula fragilis + >. + F 
Jeffreys, 1897 

Silicula filatovae + + + + d à à 
Allen & Sanders, 1973 

Silicula mcalesteri + + 
Allen & Sanders, 1973 

Malletia abyssorum d t d d + а d 
Verrill & Bush, 1898 

Malletia cuneata d + + + + 
Jeffreys, 1876 

Malletia grasslei + + 
Sanders & Allen, 1985 

Malletia johnsoni + d d d d + 
Clarke, 1961 

Malletia malita + 
Sanders & Allen, 1985 

Malletia obtusa 2 
G1O,Sars, 1878 

Malletia pallida d d d d + 
Smith, 1885 

Malletia polita d d d at 
Verrill & Bush, 1898 

Malletia surinamensis + 
Sanders & Allen, 1985 

Malletia sp. + 

Bentharca asperula d d + + + + 
(Dall, 1881) 

Bentharca nodulosa + 
(Muller, 1776) 

Bathyarca glacialis + 
(Gray, 1824) 

Bathyarca inaequisculpta + d + + $ d + 
(Smith, 1885) 

Bathyarca pectunculoides d d + + 
(Scacchi, 1834) 

Bathyarca sp. 245 + 


(continues) 


72 ALLEN 
(continued) 


ARG BRA SUR CAV NAMWEU CAN SLE GU1 ANG CAP 


Limopsis aurita + d 
(Brocchi, 1814) 
Limopsis cristata affinis d d 
Verrill, 1885 
Limopsis cristata cristata d + 
Jeffreys, 1876 
Limopsis cristata lanceolata 2 + 
Oliver & Allen, 1980 
Limopsis cristata intermedia + + + 
Oliver & Allen, 1980 
Limopsis galathea = d d + + d + + 
Knudsen, 1970 
Limopsis minuta + 
(Philippi, 1836) 
Limopsis spicata d 
Oliver & Allen, 1980 
Limopsis surinamensis d 
Oliver & Allen, 1980 
Limopsis tenella d a d + + d 
Jeffreys, 1876 
Limopsis sp. 197 ah 
Limopsis sp. 239 + 
Adipicola simpsoni + + 
(Marshall, 1900) 
Musculus discors + 
(Linné, 1767) 
Mytilidae sp. 115 ni 
Mytilidae sp. 330 + + 
Anomiidae sp. S29 
Dacrydium abyssorum + + а d d 
Allen, 1998 
Dacrydium angulare d 
Ockelmann, 1983 
Dacrydium hedleyi + 
Allen, 1998 
Dacrydium ockelmanni * + d d d d 
Mattson & Waren, 1977 
Dacrydium sandersi d + d d 
Allen, 1998 
Dacydium vitreum + + 
(Moller, 1842) 
Dacrydium wareni + = + 
Salas & Gofas, 1997 
Dacrydium sp. 2 + 
Limatula bisecta + 
Allen, 2004 
Limatula celtica + + + 
Allen, 2004 
Limatula laminifera d 
(Smith, 1885) 
Limatula louiseae + + + + + a 
Clarke, 1974 


— 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 13 
(continued) 


ARG BRA SUR CAV NAM WEU CAN SLE GU1 ANG CAP 


Limatula margaretae + + + 
Allen, 2004 
Limatula smithi + + 
Allen, 2004 
Limatula subovata d + + + + 
(Jeffreys, 1876) 
Limatula sp. 3 + 
Limea argentineae E 
Allen, 2004 
Limea lirata + 
Allen, 2004 
Limea sarsi d 
Lovén, 1846 
Limea sp. 240 + 
Flexipecten proteus + 
(Solander, 1817) 
Zygochlamys patagonica + 
(King, 1831) 
Pectinid sp. at 
Pectinidae sp. a + 
Pectinidae sp. b at + 
Pectinidae sp. c + 
Pectinidae sp. d of 
Pectinid sp. e + 
Placopecten magellanicus + 
(Gmelin, 1791) 
Similipecten minor + 
Locard, 1898 
Similipecten similis + 
(Laskey, 1811) 
Similipecten sp. + 
Delectopecten vitreus + d 
(Gmelin, 1791) 
Delectopecten sp. a d 
Hyalopecten parvulinus + 
(Locard, 1897) 
Hyalopecten undatus + + 
Verrill & Smith, 1885 
Hyalopecten sp. a + 
Hyalopecten sp. e + 
Parvamussium lucidum d + 
(Jeffreys, 1873) 
Parvamussium obliquum + 
Smith, 1885 
Parvamussium permirum + 
(Dautzenberg, 1925) 
Parvamussium sp. a + + + + + 
Parvamussium sp. b it 
Parvamussium sp. q + + 
Parvamussium sp. 2 + 
Bathypecten eucymatus + + + + 
(Dall, 1898) 


(continues) 


74 ALLEN 


(continued) 


ARG BRA SUR CAV NAMWEU CAN SLE GU1 ANG CAP 


Lee LIU nn 


Bathypecten sp. a d + ar 

Bathypecten sp. b + + 

Bathypecten sp. с + 

Bathypecten sp. d + 

Bathypecten sp. e de + 

Propeamussium centobi + 
Schein, 1989 

Propeamussium meridionale a 
(Smith, 1885) 

Propeamussium thalassinum + 
(Dall, 1886) 

Propeamussium sp. a + 

Propeamussium sp. с E 

Propeamussium sp. d + 

Pseudamussium clavatum a 
(Poli, 1795) 

Cyclopecten ambiannulatus + 
Schein, 1989 

Cyclopecten pustulosus + a 
(Verrill, 1873) 

Cyclopecten simplex u 
Verrill, 1897 

Cyclopecten sp. a d + + d + 

Cyclopecten sp. ze + i + 

Cyclopecten sp. zf + 

Cyclopecten sp. zg d + 

Lucinoma Поза + + 
(Stimpson, 1851) 

Myrtea lens + 
(Verrill 8 Smith, 1881) 

Lucinidae sp. 280 + 

Axinus grandis ab E 
(Verrill & Smith, 1885) 

Leptaxinus incrassatus d 
(Jeffreys, 1876) 

Thyasira alleni 4 + + d 
Carrozza, 1981 

Thyasira atlantica + 
Payne & Allen, 1991 

Thyasira biscayensis + # ы 
Payne & Allen, 1991 

Thyasira brevis d + d + + d 
(Verrill & Bush, 1898) 

Thyasira bushae № 
Payne & Allen, 1991 

Thyasira carrozae d + + d d 
Payne & Allen, 1991 

Thyasira croulinensis + + + d d * + 
(Jeffreys, 1847) 

Thyasira equalis a + + + + + + + + 
(Verrill & Bush, 1898) 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 15 


(continued) 


ARG BRA SUR CAV NAM WEU CAN SLE GU1 ANG CAP 


Thyasira eumyaria + d + + + + 
(M. Sars, 1870) 

Thyasira excavata plicata + 
(Verrill, 1885) 

Thyasira ferruginea d + d d d d + 
(Locard, 1886) 

Thyasira inflata d d e d + 
Payne & Allen, 1991 

Thyasira obsoleta + d + + + 
(Verrill & Bush, 1898) 

Thyasira pygmaea d A d + + 
(Verrill & Bush, 1898) 

Thyasira subcircularis Pr + “a 
Payne & Allen, 1991 

Thyasira subequatoria + à 
Payne & Allen, 1991 

Thyasira subovata minuta + + 
Payne & Allen, 1991 

Thyasira subovata subovata + + + d + + + 
(Jeffreys, 1881) 

Thyasira succisa succisa d 
(Jeffreys, 1876) 

Thyasira succisa atlantica + + + + + d + 
Payne & Allen, 1991 

Thyasira transversa d + d + d + + d 
Payne & Allen, 1991 

Thyasira trisinuta + 
(d’Orbigny, 1846) 

Thyasira tortuosa + + at + d + 
(Jeffreys, 1881) 

Thyasira ultima + + d # d 
Payne & Allen, 1991 

Thyasira verrilli + d 
Payne & Allen, 1991 

Thyasira sp. | + 

Thyasira sp. h + 

Thyasira sp. p + 

Thyasira sp. r + 

Thyasira sp. 1 

Thyasira sp. 2 a d 

Thyasira sp. 2 с 

Thyasira sp. 3 

Thyasira sp. 4 

Thyasira sp. 8 a + + 

Thyasira sp. 9 

Thyasira sp. 15 + + 

Thyasira sp. 17 af mm + 

Thyasira sp. 21 d 

Thyasira sp. 28 + + 

Thyasira sp. 30 + + + 

Thyasira sp. 32 + 


+ + + + 
+ + + + 


+ + + + + 
-- 


о + + + 
Le 
+ 


(continues) 


76 ALLEN 
(continued) 


ARG BRA SUR CAV NAMWEU CAN SLE GU1 ANG CAP 


Thyasira sp. 34 di 
Thyasira sp. 40 + + 
Thyasira sp. 45 + + + + 
Thyasira sp. 47 a + 2+ 
Thyasira sp. 47 b + 
Thyasira sp. 65 + 
Thyasira sp. 67 + 
Thyasira sp. 72 + 
Thyasira sp. 98 
Thyasira sp. 128 + 
Thyasira sp. 186 
Thyasira sp. 188 
Thyasira sp. 188 b 
Thyasira sp. 202 
Thyasira sp. 210 + 
Thyasira sp. 239 + 
Thyasira sp. 243 
Thyasira sp. 280 + 
Thyasira sp. 297 + 
Thyasira sp. 299 
Thyasira sp. 306 + 
Thyasira sp. 309 
Thyasira sp. 314 
Thyasira sp. 318 
Thyasira sp. 323 
Thyasira sp. 328 
Thyasira sp. 329 + 
Thyasira sp. 330 + 
Thyasira sp. 334 
Thyasira sp. 346 a 
Thyasira sp. 346 b 
Thyasira sp. 346 c 
Thyasira sp. DS87 + 
Thyasira sp. S50 + 
Thyasira sp. 6797 + 
Galeommatoidea sp. + 
Epilepton elpis + 
Allen, 2007 
Epilepton subtrigonum + 
(Fischer, 1873) 
Epilepton sp. 21 + + + 
Leptonidae sp. v + 
Leptonidae sp. w + 
Montacuta ovata oh + + à 
Jeffreys, 1881 
Montacuta sp. + 
Montacuta sp. 1 + + 
Axinodon symmetros + + т 
(Jeffreys, 1876) 
Modiolarca tumida + 
(Hanley, 1843) 


+ 


+ + + + 


En 


+ 


+ + + + + 


+ + + + 


(continues) 


(continued) 


Mysella tumidula 
(Jeffreys, 1866) 
Mysella verrilli 
(Dall, 1899) 
Mysella sp. 1 
Mysella sp. 2 
Mysella sp. 3 
Neolepton profundorum 
Allen, 2000 
Abra longicallis 
(Scacchi, 1834) 
Abra profundorum 
(Smill, 1885) 
Tellinidae sp. a 
Tellinidae sp. b 
Tellinidae sp. c 
Tellinidae sp. d 
Tellinid sp. f 
Tellinid sp. x 
Kelliella abyssicola 
Allen, 2001 
Kelliella adamsi 
(Smith, 1885) 
Kelliella atlantica 
(Smith, 1885) 
Kelliella biscayensis 
Allen, 2001 
Kelliella concentrica 
Allen, 2001 
Kelliella elongata 
Allen, 2001 
Kelliella miliaris 
Philippi, 1844 
Kelliella nitida 
Verrill, 1885 
Kelliella tenina 
Allen, 2001 
Kelliella sp. 236 
Kelliella sp. 245 
Kelliella sp. 323 
Carditidae sp. 1 
Carditidae sp. 2 
Carditidae sp. 5 
Goodallia triangularis 
(Montagu, 1803) 
Astarte sp. 1 
Astarte sp. 2 
Astarte sp. 5 
Tridonta elliptica 
(Brown, 1827) 
Acanthocardia echinata 
(Linne, 1758) 


BIVALVIA OF THE DEEP ATLANTIC 


ra 


ARG BRA SUR CAV NAMWEU CAN SLE GU1 ANG CAP 


+ 
+ + + 
+ + + + 
+ + 
d 
+ + 
+ + + 
+ 
+ 
+ 
+ + 
+ 
d d d HE d 
d 
2. 
+ + + 
+ 
+ 
+ 
+ 
+ 
+ 
d 
+ 
+ 
d + 
d 
+ 
+ 
+ 


+ 


(continues) 


78 ALLEN 


(continued) 


ARG BRA SUR CAV NAMWEU CAN SLE GU1 ANG CAP 


Clinocardium ciliatum 5 
(O. Fabricius, 1780) 
Cardiidae sp. 4 + 
Cardiidae sp. 6 
Veneridae sp. 280 a 
Veneridae sp. h = 
Mactridae sp. + 
Corbula sp. 1 + 
Corbula sp. 2 + 
Lyonsia sp. 1 + 
Pandora pinna + 
(Montagu, 1803) 
Asthenotherus hemphilli + 
Dall, 1886 
Thracia conrad! + 
Couthouy, 1838 
Thracia durouchouxi + 
Dautzenberg € Fischer, 1897 
Thracia gracilis + 
Jeffreys, 1865 
Thracia nitida ge + + 
Verrill, 1884 
Thracia myopsis h 
(Maller, 1842) 
Thracia pubescens + 
(Pulteney, 1799) 
Thracia sp. 1 $ 
Thracia sp. 2 As 
Thracia sp. 3 FE ES 
Periploma papyracea ir 
(Poli, 1791) 
Bushia sp. + 
Cochlodesma tenerum + + 
(Jeffreys, 1880) 
Xylophaga sp. + 
Poromya granulata A + 
(Nyst 8 Westendorp, 1839) 
Poromya tornata + + LA 
(Jeffreys. 1876) 
Poromya sp. 256 + 
Poromya sp. 301 + 
Роготуа sp. 335 + 
Cetoconcha angolensis + 
Allen & Morgan, 1981 
Cetoconcha braziliensis + 
Allen & Morgan, 1981 
Cetoconcha sp. + 
Protocuspidaria atlantica + + 
Allen & Morgan, 1981 
Protocuspidaria simplis + + + + 
Allen & Morgan, 1981 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 79 
(continued) 


ARG BRA SUR CAV МАМ WEU CAN SLE GU1 ANG CAP 


Protocuspidaria verityi Е à =; = + 5 + 
Allen & Morgan 1981 
Protocuspidaria sp. 256 nz 
Protocuspidaria sp. + 
Cuspidaria atlantica + + + + + 
Allen & Morgan, 1981 
Cuspidaria barnardi + + ch 
Knudsen, 1970 
Cuspidaria circinata + + 
(Jeffreys, 1876) 
Cuspidaria jeffreysi + + + 
(Dall, 1881) 
Cuspidaria obesa + d d 
(Loven, 1846) 
Cuspidaria parva + + + + + + + 
Verrill & Bush, 1898 
Cuspidaria undata + 
(Verrill, 1884) 
Cuspidaria ventricosa + 
Verrill & Bush, 1898 
Cuspidaria sp. + 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 
Cuspidaria sp. 56 7 
Cuspidaria sp. 126 + 
Cuspidaria sp. 202 + 
Cuspidaria sp. 239 + 
Cuspidaria sp. 242 + 
Cuspidaria sp. 326 + 
Cuspidaria sp. 328 + 
Cuspidaria sp. 334 + 
Cuspidaria sp. 346 + 
Cuspidaria sp. 519 + 
Cardiomya costellata + + + 
(Deshayes, 1830) 
Cardiomya curta + 
(Jeffreys, 1876) 
Cardiomya knudseni + + + + 
Allen & Morgan, 1981 


CON Боро ооо 
+ © + + + 
+ 


(continues) 


80 ALLEN 
(continued) 


ARG BRA SUR CAV NAMWEU CAN SLE GU1 ANG CAP 


Rhinoclama abrupta + + + 
Allen & Morgan, 1981 

Rhinoclama halimera + + + + 
(Dall, 1836) 

Rhinoclama notabilis d + + + 
(Jeffreys, 1876) 

Luzonia simplex d d d d 
Allen & Morgan, 1981 

Tropidomya abbreviata + + + + 
(Forbes, 1843) 

Tropidomya diagonalis + 
Allen & Morgan, 1981 

Halonympha atlanta + + + 
Allen 8 Morgan, 1981 

Halonympha depressa + # + 
(Jeffreys, 1881) 

Myonera angularis + 
(Jeffreys, 1876) 

Myonera atlantica + + + + d 
Allen & Morgan, 1981 

Myonera demistriata + + + + 
Allen & Morgan, 1981 

Myonera octoporosa au + 
Allen & Morgan, 1981 

Myonera paucistriata d + + + 
Dall, 1886 

Myonera tillamookensis x 
Dall, 1916 

Myonera sp. d + 

Lyonsiella abyssicola + + + # + 
(С. О. Sars, 1872) 

Lyonsiella formosa + + + + + 
(Jeffreys, 1881) 

Lyonsiella fragilis + 
Allen & Turner, 1974 

Lyonsiella freilei + + 
Allen & Turner, 1974 

Lyonsiella perplexa + + 
Allen & Turner, 1974 

Lyonsiella smidti ce + + 
(Smith, 1885) 

Lyonsiella subquadrata # + + 
(Jeffreys, 1881) 

Lyonsiella sp. 202 + 

Lyonsiella sp. 239 di 

Verticordia quadrata A + + + + + * 
Smith, 1885 

Verticordia triangularis + + + + + 
Locard, 1898 

Verticordia sp. + 

Verticordia sp. 88 + 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 81 
(continued) 


ARG BRA SUR CAV NAMWEU CAN SLE GU1 ANG CAP 


Verticordia sp. 98 + 
Verticordia sp. 240 + 
Verticordia sp. 309 + 
Policordia atlantica + + + + 

Allen & Turner, 1974 
Policordia densicostata + + + 

| осага, 1898 
Policordia gemma + + + + + 

(Verrill, 1880) 
Policordia insoleta + + + 

Allen & Turner, 1974 
Policordia laevis À + 

Allen & Turner, 1974 
Polycordia jeffreysi + + 

(Friele, 1879) 
Policordia sp. 128 + 
Policordia sp. 200 + 
Policordia sp. 297 + 
Laevicordia horrida + + в 

Allen & Turner, 1974 
Laevicordia sp. + 
Halicardia flexuosa + 

(Verrill & Smith, 1881) 
Incertae cedis зр. 1 а + + 
Incertae cedis sp. 1 b + 
Incertae cedis sp. 2 + 
Incertae cedis sp. 3 
Incertae cedis sp. 10 + 
Incertae cedis sp. 64 
Incertae cedis sp. 72 
Incertae cedis sp. 89 
Incertae cedis sp. 95 
Incertae cedis sp. 96 
Incertae cedis sp. 100 
Incertae cedis sp. 101 + 
Incertae cedis sp. 115 
Incertae cedis sp. 119 
Incertae cedis sp. 121 
Incertae cedis sp. 122 
Incertae cedis sp. 131 
Incertae cedis sp. 146 + 
Incertae cedis sp. 185 
Incertae cedis sp. 186 + 
Incertae cedis sp. 188 
Incertae cedis sp. 189 
Incertae cedis sp. 202 a 
Incertae cedis sp. 202 b 
Incertae cedis sp. 203 a $ 
Incertae cedis sp. 203 b + 
Incertae cedis sp. 236 + 
Incertae cedis sp. 237 + 


+ +++ ++++++ + 
= 


+++ + 


(continues) 


82 ALLEN 


(continued) 
ARG BRA SUR CAV NAMWEU CAN SLE GU1 ANG CAP 
Incertae cedis sp. 240 + 
Incertae cedis sp. 256 + 
Incertae cedis sp. 262 + + + 


Incertae cedis sp. 291 a 
Incertae cedis sp. 291 b 
Incertae cedis sp. 299 
Incertae cedis sp. 301 
Incertae cedis sp. 313 
Incertae cedis sp. 314 + 
Incertae cedis sp. 318 

Incertae cedis sp. 323 

Incertae cedis sp. 326 

Incertae cedis sp. 328 

Incertae cedis sp. 330 + + 
Incertae cedis sp. 334 a 

Incertae cedis sp. 334 b 

Incertae cedis sp. 335 

Incertae cedis sp. 346 


+ + + + 


++ + + + + + + 


+ + + + + 
+ 


APPENDIX 2 


The depth distribution of first, second and third dominant deep-sea species of bivalves in differ- 
ent basins giving minimum (min) and maximum (max) depth range and depth of maximum 
abundance (opt). + indicates the species is dominant at one station in a particular basin and a 
figure in paraentheses indicates that a species is dominant at more than one station in the 
basin. 


Dominance Depth (m) 

Species 1 2 3 Basin min max Opt 
Pristigloma nitens + WEU 2,397 4,734 3,338 
Deminucula atacellana + МЕО 3,919 4,400 4,400 
+) + МАМ 530 *.3.9384+ 02022 
+(3) АКС ПО пе SONG 3680 
+ (2) WEU 1,500 2,493 4,286 
Brevinucula verrilli + САМ 2,129. 2988. u 72,929 
+ SLE 1,976 3,861 » 22,934 
+ GUI 2,470 3,174 2,470 
Microgloma turnerae + WEU MO 3 2.398922 
Microgloma turnerae + + SUR 2076 ERRORS 
+ ANG 2.031 2154 2,784 
Microgloma sp. $ + + SUR 1,022 2289951810 
Phaseolus sp. с + ARG 5223 1 6223 15223 
Nuculoidea bushae + NAM 196 530 530 
+ WEU 860 2,493 1,015 
+ CAN 780: 1 2,129 A229 

mi GUI 1201.20 29610921161 
Nuculoidea pernambucensis + BRA 827 1007 827 
Nuculoma granulosa +(2) + NAM 390.100 И =. 102 
u WEU 1/3360" 1390 1336 


(continues) 


(continued) 


Species 


Nuculoma perforata 
Nuculoma similis 
Tindaria agatheda 
Tindaria miniscula 
Tindaria callistiformis 
Neilonella salicensis 


Neilonella whoii 


Pseudotindaria erebus 


Nuculana vestita 
Propoleda carpenteri 
Prelametila clarkei 


Lametila abyssorum 
Ledella acinula 
Ledella acuminata 
Ledella jamesi 


Ledella pustulosa argentinea 
Ledella pustulosa marshalli 
Ledella pustulosa pustulosa 


Ledella sublevis 


Ledella ultima 


Portlandia minuta 
Spinula filatovae 


Spinula subexisa 
Yoldiella americana 


Yoldiella bilanta 
Yoldiella biscayensis 
Yoldiella blanda 
Yoldiella capensis 
Yoldiella curta 


BIVALVIA OF THE DEEP ATLANTIC 


Dominance 
1 2 3 
+(3) 
= + + 
+ 

+ 
+ 
+(3) 
+ + 
a2) KZ) 
+ 
e 
+ +(2) 
+ +(8) 
+ 
++ 
+ 
+ 
+ 
+ 
+ 
+e 
+ 
+(2) 
+ -- 
+ +(2) 
+ +(3) 
AN Zi) 
+ 
+ 
+ 
+(12) +(2) 
+ 
+(2) 
+(2) + + 
+ + 
+(4) 
4) 
nu 
+ 
+ 
+ 
+ +(6) 
+(2) 
+(2) 
+ +(3) + 
+(2) 
+ 
+(4) + 
+ 
a 
+ +(3) 


Basin 


SEE 
NAM 
CAN 
NAM 
NAM 
NAM 
SUR 
WEU 
GUI 
ANG 
NAM 
WEU 
ANG 
ANG 
ANG 
ARG 
NAM 
ARG 
BRA 
SUR 
WEU 
SUR 
ARG 
ARG 
WEU 
WEU 
ARG 
GUI 
NFD 
NAM 
SUR 
CVD 
WEU 
SEE 
ANG 
CAP 
ANG 
GUI 
ANG 
GUI 
NAM 
SUR 
CAP 
WEU 
ARG 
CAP 
NAM 
SUR 
BRA 
WEU 
ANG 


Depth (m) 
min max 
1,796 ‘255 
475 475 
19384 2.129 
S53 Corse 
3,153 ° - 5042 
1,102 2,886 
523 2076 
641 2,503 
2470 2,514 
ар 2081 
2,862 4,970 
3,338 4,823 
2754 4,596 
2 MODA IBAS 
542 ı 2035 
1041 2 ЗАЗ 
2,886 2,886 
ДОР whe225 
3,495: 3,495 
1,022 2076 
2540; «> 2900 
1.513 2.2353 
1,679 2,480 
Solr 9290 
2,494. 4.425 
609 3,859 
WORD: ER 
2470 DB 514 
3,919 4,400 
2,900 5,042 
3099 5150 
5867 5,673 
3.338 483 
2,891 3.028 
2,031 4,630 
4,560 5280 
542 542 
1,261 1,261 
1,643 1,643 
1,261 1,261 
3,834 5,042 
4429 5150 
622 2,864 
2111 4794 
3910 52 
622 1,014 
811 2,176 
1,022 - och Ste 
827 1,007 
1,500: 92,808 
2,992 3,779 


83 


(continues) 


84 


(continued) 


Species 


Yoldiella ella 


Yoldiella enata 
Yoldiella extensa 
Yoldiella frigida 


Yoldiella jeffreysi 


Yoldiella inconspicua inconspicua 


Yoldiella inconspicua africana 


Yoldiella insculpta 
Yoldiella lata 

Yoldiella lucida 
Yoldiella obesa incala 
Yoldiella ovata 
Yoldiella perplexa 
Yoldiella pseudolata 
Yoldiella robusta 


Yoldiella subcircularis 
Yoldiella subequilateria 


Silicula filatovae 
Malletia abyssorum 


Malletia cuneata 


Malletia johnsoni 


Malletia pallida 


Malletia polita 


Bentharca asperula 


Bathyarca inaequisculpta 


+(2) 


ALLEN 
Dominance 
2 5) 
+ 
+ +(6) 
== 
Ah 
+ 
+(2) 
+ + 
A A) 
+ a 
+3) #02) 
+ 
+(3) 
+(2) +(3) 
Ha а 
+ + 
+ 
+ + 
+ + 
+ 
= 
+ 
+ 
(7) HI) 
+ + 
= 
+ 
+(3) + 
+(4) +(6) 
+ 
+ + 
2) 
+ 
+ 
+ 
E 
+ 
+(2) 
fe 
+. +(2) 


Basin 


NFD 
WEU 
SLE 
SUR 
ARG 
NAM 
WEU 
NFD 
NAM 
SUR 
WEU 
NAM 
WEU 
ANG 
CAP 
WEU 
WEU 
NAM 
WEU 
SUR 
SUR 
WEU 
ARG 


NAM 
SUR 
WEU 
ANG 
NFD 
NAM 
ARG 
WEU 
SLE 
CAP 
ARG 
WEU 
NAM 
WEU 
SLE 
ANG 
NAM 
CVD 
SLE 
ANG 
NAM 
SUR 
CVD 
NAM 
ANG 
SUR 
ANG 


Depth (m) 
min max 
3,919 4,400 
3,200 4,823 
3,828 3,861 
250070923858 
ZLOTY “2707 

475 811 

609 609 
3,919 4,400 
2,064 4,862 
2,853 4,980 
2,006 4,823 

498 4,743 
1,739 4,462 
AOS’ 121091 
2,154 + 27864 
1,918 772,360 
1,865: 2,790 

475 498 
1,913 2,818 
2500 72,853 
1518 ao 2076 
1,500 2,397 

293 2323 
4,694 5,007 
4,429 5,150 
2,540 . 2,900 
2,992 4,630 
4,400 4,793 
2.864 5,042 
3,916 4,435 
2,790 4,734 
31628 3,861 
4,560 5,280 
2,480 3,916 
2,081 4,734 

478 3,834 
12739 2773 
1,790 2,934 
2.0831 3,985 
3,0 2406561007 
5873 15679 
2357 120934 
2.194 4,596 
4,750 4,750 
ой | 1516 
5,867 5,867 
2223. 4,970 
4,079 4,630 
25990 - 3.030 
2,031 4,630 


Opt 


ls 
3,480 
3,861 
2,500 
2207 

498 

609 
Soe 
2,862 
3,868 
2719 
3,356 
2,001 
2,031 
2,864 
PAS a) 
2,076 

478 
209 
2,900 
2,076 
1,500 

296 


5,007 
4,429 
2,900 
2,992 
4,400 
4,833 
4,435 
4,462 
3,901 
4,585 
2,480 
2,897 
2,496 
1992 
2,934 
3,909 
5,007 
5,9079 
2,9917 
SAUT 
SOS 
5,100 
5,867 
3,860 
4,630 
5,100 
4,566 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 85 


(continued) 


Dominance Depth (m) 

Species 1 2 3 Basin min max Opt 
Bathyarca pectunculoides . + NAM TS dr 5 
+ WEU 619 619 619 
Limopsis aurita + WEU 609 619 609 
Limopsis cristata agg. + МАМ 478-- 2.178 2,022 
+ SUR 1,91 122076: ASS 
+ WEU 1048 "16500: 14500 
Limopsis galathea +(2) + SUR 3,835 4,980 4,980 
+ BRA INES ARS OS 
+(3) + ANG 2,754 4,829 4,630 
Limopsis spicata + ARG 2,048 2,480 2,048 
Limopsis surinamensis + SUR 525 523 923 
Limopsis tenella + BRA 3,4951 8/783r ¿PES 
+(2) SLE 3,028 1.3051 Sol 
ANG 3,1971 4,596 398 
Delectopecten vitreus + WEU 485 485 485 
Delectopecten sp. a + WEU 644.7: 1,330. 13960 
Bathypecten sp. d + ARG 1,679 - . 161900 MOR 
Parvamussium lucidum + BRA 1,007 1,493 1,007 
Cyclopecten sp. a + ARG 2.3237 3,82%" 00,929 
+ WEU 3,338 4,435 4,435 
Cyclopecten sp. zg + BRA 3,495 3,495 3,495 
Limatula laminifera 12 NAM 1,1887 13,396 u 
Limatula subovata + ARG 2,107 + 2707 27 
Limea lirata + NAM 2.223 222% 12223 
Limea sarsi + WEU 619 619 619 
Dacrydium abyssorum +(9) WEU 3,992 4,823 4,228 
+ SLE 3,861 » -3:861 r «881 
+(2) CAP 4,560 5,280 4,585 
Dacrydium angulare + ANG 3,985. 3985 3,985 
Dacrydium ockelmanni + + NAM 1102, 222%. 3234 
+ WEU 1,500. 2000 2155 
+ CAN 1,934 2,129 1,934 
+ SLE 1,796 2,934 1,796 
Dacrydium sandersi +(3) + NAM 2.223: 2.000 2602 
+(2) + ВКА 1,007 3:783 3495 
+ WEU 2,775 ~ 35487 3548 
Thyasira alleni + CAP 622 1559 622 
Thyasira brevis + ARG 2,480 3,822 2,480 
+ +(2) WEU 1,739, 823 0014239 
+ CAP 2,864 2,864 2,864 
Thyasira carrozae +(2) ARG 1,079 J дом DS 
+ ANG 542 1,432 1,432 
+(2) + + CAP 481 1559 622 
Thyasira croulinensis + WEU 119 - 9,859 641 
+ SLE 2,192. 3801 ER 
Thyasira eumyaria + WEU 641 1,739 641 
Thyasira ferruginea + + NAM 196 4,825 2,064 
+ ARG 293 4,435 293 
+ WEU 485 4,466 4,466 
$: SLE 1,976 2934 208 


+ АМС 1,643 2,992 2,992 


(continues) 


86 


(continued) 


Species 


Thyasira incrassata 
Thyasira inflata 


Thyasira obsoleta 
Thyasira pygmaea 


Thyasira subovata subovata 
Thyasira succisa Succisa 
Thyasira succisa atlantica 


Thyasira tortuosa 
Thyasira transversa 


Thyasira ultima 


Thyasira verrilli 
Thyasira sp. 2a 
Thyasira sp. 17 
Thyasira sp. 21 

Carditid sp. 2 

Astarte sp. 1 

Astarte sp. 2 

Neolepton profundorum 
Kelliella atlantica 


Kelliella biscayensis 
Abra profundorum 
Tellinid sp. с 
Cuspidaria obesa 
Cuspidaria parva 


Cuspidaria sp. 1 
Cuspidaria sp. 2 
Myonera atlantica 
Myonera paucistriata 
Myonera sp. 
Rhinoclama notabilis 
Luzonia simplex 


ALLEN 
Dominance 
2 3 
+ 
+ 
+(2) 
+ 
+(2) 
+ 
+ +(2) 
42) 
+ 
+ 
+ 
+ 
+ 
+(2) 
+ 
oe 
+2) 
+(2) 
+(2) 
+ 
+ 
(0) + 28) 
+ 
+ 
== 
+(2) 
+ 
+ 


Basin 


WEU 
SUR 
ARG 
ANG 
WEU 
NAM 
ARG 
WEU 
WEU 
ANG 
ANG 
NAM 
SUR 
ARG 
WEU 
CAP 
SLE 
ANG 
NAM 
ARG 
NAM 
ARG 
WEU 
NAM 
NAM 
ARG 
SUR 
CVD 
ARG 
WEU 
CAN 
SEE 
ANG 
CAP 
WEU 
WEU 
CAP 
WEU 
NAM 
WEU 
SUR 
WEU 
CAP 
SUR 
BRA 
ARG 
WEU 
ЗЕЕ 
АМС 
САР 


Оер (т) 
min max 
ZOO ZO TS 
2.500 3,429 
3.343 "29,916 
4566 4,596 
860 2,868 
Ato, 70 
2323 9° 2323 
2:081 45254 
619 1,950 
ое SGA 
1432. - 1,643 
3806 “46893 
523 5100 
1,011 5.223 
860 860 
622 52 2154 
1,796: 2357 
14.643: 72,081 
2.0644 32223 
293 293 
AND 103.854 
4402. 4.402 
119 485 
119 530 
475 498 
Ро 27.07 
523 4.980 
5,867 5.867 
2,480 4,402 
1500 474 
1,934 2,988 
11976. 2557 
542 4,596 
481 5,240 
ASS “12015 
1015 45828 
481 481 
2,081 3,548 
478 3,806 
609 4,632 
3.968; 15100 
619 619 
A560 4/685 
1.022, 14022 
416 1,493 
1:04 1,011 
1,894, 2430 
11196. РЭ 
mae. 41091 
622. 2194 


Opt 


2,928 
3,429 
3,343 

096 
1,739 

530 
2329 
2,494 
1,336 
1,643 
1,432 
4,833 
3,100 
2,048 

860 

622 
ZZ 
1,643 
2,228 

299 
14102 
4,402 

485 

478 

478 
O 

20 
5,867 
3,343 
2505 
1,934 
2301 
2,031 

481 
AOS 
1,015 

481 
2920 
1,254 
2130 
3,868 

619 
4,560 
1,022 
1,007 
1,011 
1,894 
17976 

942 

329 


BIVALVIA OF THE DEEP ATLANTIC 
APPENDIX 3 


List of living bivalves taken in epibenthic sledge samples from various 
deep-sea expeditions in the Atlantic arranged in depth sequence in each 
individual basin. The number of individuals and their percentage occur- 
rence in each sample is given. 


No. of Occurrence 
Sample Individuals in % 


NEWFOUNDLAND BASIN 
Sta. 335, 3,919 т, 40°42.6’N, 46°30.0’W 


Deminucula atacellana 3 28 
Neilonella whoii 5 4.1 
Ledella ultima 22 18.2 
Yoldiella ella 26 2:8 
Yoldiella jeffreysi 28 23.1 
Yoldiella subcircularis 2 12 
Limopsis galathea 5 4.1 
Propeamussium sp. a 1 0.8 
Thyasira ferruginea 1 0.8 
Thyasira transversa IR 14.1 
Abra profundorum 2 17 
Poromya sp. 335 1 0.8 
Protocuspidaria verityi 1 0.8 
Cuspidaria parva 1 0.8 
Cuspidaria Sp. | 1 0.8 
Myonera atlantica 2 Ah 
Rhinoclama notabilis 2 27 
Incerte cedis sp. 335 1 0.8 
Sta. 334, 4,400 т, 40°42.6’М, 46°13.8’W 
Pristigloma alba 1 0.1 
Pristigloma nitens © 0.3 
Microgloma pusilla 17 8 
Deminucula atacellana 173 17.8 
Neilonella whoii 3 Ome! 
Ledella ultima 121 129 
Yoldiella americana 102 DOS 
Yoldiella dissimilis 28 2.4 
Yoldiella ella 76 1.8 
Yoldiella enata 6 0.6 
Yoldiella inconspicua inconspicua 28 2.9 
Yoldiella jeffreysi 80 8.3 
Yoldiella subcircularis 8 0.8 
Silicula filatovae 2 Or 
Malletia abyssorum 121 125 
Malletia cuneata 9 0.9 
Malletia polita 7 OT 
Hyalopecten sp. a 6 0.6 
Parvamussium Sp. Z 10 1.0 
Bathypecten eucymatus 4 0.4 
Thyasira brevis 6 0.6 
Thyasira equalis 1 0.4 


(continues) 


87 


ALLEN 


(continued) 


Sample 


Thyasira transversa 
Thyasira sp. 15 
Thyasira sp. 334 
Montacuta sp. 1 
Mysella verrilli 

Abra profundorum 
Poromya sp. 335 
Protocuspidaria verityi 
Cuspidaria inflata 
Cuspidaria parva 
Cuspidaria sp. 334 
Halonympha atlanta 
Myonera atlantica 
Incerte cedis sp. 3 
Incerte cedis sp. 314 
Incerte cedis sp. 334 a 
Incerte cedis sp. 334 b 


Sta. 331, 4,793 m, 41°13.0’N, 41°36.7’W 


Pristigloma nitens 
Yoldiella americana 
Yoldiella subcircularis 
Malletia abyssorum 
Malletia polita 
Dacrydium abyssorum 
Limatula louiseae 
Limatula sp. 3 
Thyasira transversa 
Astarte triangularis 
Abra profundorum 
Rhinoclama notabilis 


NORTH AMERICA BASIN 


Sta. 170, 68 т, 40°37.0’, 70°50.0’W 
Placopecten magellanicus 
Periploma papyracea 
Lucinoma filosa 


Sta. 172, 119 m, 40°12.3’N, 70°44.7’W 
Nuculana acuta 
Thyasira trisinuta 
Carditidae sp. 1 
Astarte sp. 1 


Sta. 89, 196 m, 40%01.6'N, 70°40.7’W 
Nuculoidea bushae 
Nuculana acuta 
Malletia sp. 
Bathyarca pectunculoides 
Dacrydium vitreum 
Thyasira croulinensis 
Thyasira ferruginea 
Thyasira obsoleta 


No. of 
Individuals 


32 


— 


© O1 = N фм = © © — 


72 
286 


— 


= NN 


Occurrence 
in % 


39 
0.9 


o0000090-2000% 
N = N = © O1 © © © © = = © 


(continues) 


(continued) 
No. of 
Sample Individuals 

Mysella sp. 1 1 
Mysella sp. 2 2 
Cuspidaria sp. f 4 
Cardiomya knudseni 7 

Sta. 346, 475 m, 39°54.1’N, 70°10.7’W 
Nuculoidea bushae 4 
Nuculoma similis 205 
Yoldiella frigida 43 
Yoldiella lucida 285 
Dacrydium vitreum 4 
Cyclopecten pustulosus 5 
Thyasira croulinensis 3 
Thyasira ferruginea 17 
Thyasira obsoleta 1 
Thyasira pygmaea 236 
Thyasira succisa atlantica 8 
Thyasira transversa 1 
Thyasira sp. 15 16 
Thyasira sp. 17 1 
Thyasira sp. 346 a 1 
Thyasira sp. 346 b 1 
Thyasira sp. 346 c 2 
Astarte Sp. 2 218 
Abra longicallis 4 
Thracia conradi 1 
Laevicardia horrida 24 
Cuspidaria parva 27 
Cuspidaria sp. a 4 
Cuspidaria sp. k & 
Cuspidaria sp. 346 6 
Lyonsiella abyssicola 3 
Lyonsiella perplexa 10 
Incerte cedis sp. 346 4 

Sta. 88, 478 m, 39°54.1’N, 70°37.0’W 
Nuculoma similis 35 
Nuculana acuta 1 
Yoldiella lucida 288 
Malletia johnsoni 1 
Bathyarca glacialis 1 
Bathyarca pectunculoides 3 
Limopsis cristata agg. 107 
Dacrydium vitreum 34 
Cyclopecten pustulosus 3 
Thyasira croulinensis 28 
Thyasira equalis 5 
Thyasira ferruginea 41 
Thyasira pygmaea 26 
Thyasira sp. 17 2 
Astarte sp. 1 108 
Astarte sp. 2 ED 


BIVALVIA OF THE DEEP ATLANTIC 


Occurrence 
in % 


0:9 
1.8 
3.6 
6.3 


= O) 
wOoW=200050w 
© © OO NN © A HW = 


(continues) 


89 


ALLEN 


(continued) 


No. of Occurrence 


Sample Individuals in % 
Kelliella atlantica 1 0.1 
Kelliella elongata 3 0.3 
Thracia nitida 1 0.1 
Verticordia sp. 88 2 0.2 
Laevicordia horrida 1 0.1 
Cuspidaria atlantica 2 0.2 
Cuspidaria parva 73 1.7 
Lyonsiella abyssicola 14 4:5 
Incerte cedis 1 0.1 
Sta. 96, 498 m, 39°55.2’N, 70°39.5’W 
Nuculoma similis 86 10.9 
Yoldiella frigida 120 15.3 
Yoldiella inconspicua inconspicua 1 0.1 
Yoldiella lucida 185 29.5 
Bathyarca pectunculoides 1 0.1 
Limopsis aurita 2 0.3 
Limopsis cristata agg. 60 7.6 
Thyasira croulinensis 14 1:8 
Thyasira ferruginea 45 5:7 
Thyasira pygmaea 62 7.9 
Mysella sp. 1 1 0.1 
Mysella sp. 2 1 0.1 
Kelliella elongata 2 0.3 
Astarte sp. 2 138 17.5 
Cuspidaria atlantica 3 0.4 
Cuspidaria parva 3g 3.0 
Cuspidaria sp. (broken) 1 0.1 
Lyonsiella abyssicola 21 2:5 
Sta. 105, 530 m, 39°56.6’М, 71°03.6’W 
Deminucula atacellana 6 0.4 
Nuculoidea bushae 259 190 
Nuculoma granulosa 232 14.8 
Yoldiella frigida 129 8.2 
Yoldiella lucida 126 8.1 
Malletia johnsoni 3 0.2 
Bathyarca pectunculoides 3 0.2 
Thyasira croulinensis on 2.4 
Thyasira equalis alé 7.4 
Thyasira pygmaea 459 29.2 
Thyasira subovata subovata 1 0.06 
Astarte sp. 1 149 9.5 
Kelliella concentrica 5 0.3 
Lyonsia sp. 1 1 0.06 
Cuspidaria atlantica 1 0.06 
Cuspidaria parva 51 3.9 
Lyonsiella abyssicola 19 eZ 
Sta 207.011 m, oo 51.3N, 70°54. 3° VV 
Solemya acherax 7 0.2 
Nuculoma granulosa 61 1.4 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Nuculoma similis 
Yoldiella curta 

Yoldiella frigida 
Yoldiella lucida 
Bathyarca pectunculoides 
Limopsis cristata affinis 
Limatula subovata 
Thyasira equalis 
Thyasira ferruginea 
Thyasira obsoleta 
Thyasira pygmaea 
Thyasira tortuosa 
Thyasira sp. 17 
Thyasira sp. 45 

Kelliella concentrica 
Cuspidaria sp. f 

Sta. 87, 1,102 m, 39°48.7’N, 70°40.8’W 
Solemya grandis 
Deminucula atacellana 
Nuculoma granulosa 
Nuculoma similis 
Neilonella salicensis 
Yoldiella curta 

Yoldiella inconspicua inconspicua 
Limopsis cristata affinis 
Dacrydium ockelmanni 
Limatula subovata 
Bathypecten eucymatus 
Thyasira croulinensis 
Thyasira equalis 
Thyasira ferruginea 
Thyasira obsoleta 
Thyasira pygmaea 
Thyasira subovata subovata 
Thyasira tortuosa 
Thyasira sp. 1 

Thyasira sp. 2 а 
Thyasira sp. 15 
Thyasira sp. 17 

Thracia conradi 
Policordia densicostata 
Cuspidaria obesa 
Cuspidaria parva 
Lyonsiella abyssicola 
Lyonsiella fragilis 


Sta. 118, 1,153 т, 32°19.4’N, 64°34.9'W 


Neilonella salicensis 
Phaseolus sp. d 
Yoldiella enata 
Malletia johnsoni 


No. of 
Individuals 


1348 
259 


31 


Occurrence 
in % 


31.6 
5.6 
3:6 
6.2 


(continues) 


91 


ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Bathyarca pectunculoides 24 15:2 
Dacrydium sandersi 5 32 
Dacrydium wareni 1 0.6 
Limatula laminifera 26 16.5 
Parvamussium sp. q 10 6.3 
Cyclopecten simplex 1 0.6 
Cyclopecten sp. ze 2 1.3 
Thyasira equalis 4 2.5 
Thyasira succisa atlantica 7 4.4 
Thyasira sp. 1 1 0.6 
Thyasira sp.2 a 2 £3 
Thyasira sp. 17 1 0.6 
Thyasira sp. 11 8.2 
Corbula sp. 1 1 0.6 
Protocuspidaria atlantica 1 0.6 
Protocuspidaria verity! 1 0.6 
Cuspidaria rostrata 1 0.6 
Cardiomya curta 1 0.6 
Myonera atlantica 14 8.2 
Myonera demistriata 1 0.6 
Lyonsiella abyssicola 2 ques 

Sta. 73, 1,470 т, 39°46.5’М, 70°43.3’W 
Solemya grandis 2 0.04 
Deminucula atacellana 361 7.9 
Nuculoma granulosa 1143 25:0 
Nuculoma similis 270 5,9 
Neilonella salicensis 497 10.9 
Ledella parva 1 0.02 
Yoldiella curta 699 158 
Yoldiella inconspicua inconspicua 1 0.02 
Malletia johnsoni 341 7.5 
Limopsis cristata affinis 297 6.5 
Dacrydium ockelmanni 90 2.0 
Dacrydium wareni 2 0.04 
Limatula laminifera 1 0.02 
Limatula subovata Я 2.4 
Bathypecten eucymatus 100 22 
Thyasira croulinensis 40 0.9 
Thyasira equalis 5 0.1 
Thyasira ferruginea 136 3.0 
Thyasira pygmaea 5 0.1 
Thyasira tortuosa 4 0.09 
Thyasira sp. 15 40 0.9 
Thyasira sp. 17 362 7.9 
Policordia densicostata 1 0.02 
Cuspidaria parva oy 122 
Lyonsiella abyssicola 10 0.2 
Lyonsiella fragilis 6 0.1 

Sta. 128, 1,254 m, 39°46.5’N, 70°45.2'W 
Deminucula atacellana 12 4.9 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Nuculoma granulosa 5 2.1 
Nuculoma similis ae dit 
Neilonella salicensis 12 7.4 
Yoldiella curta 8 38 
Yoldiella obesa obesa 1 4.5 
Malletia johnsoni 19 7.8 
Limatula subovata 11 4.5 
Dacrydium ockelmanni 38 15.6 
Bathypecten eucymatus 3 $2 
Thyasira croulinensis 5 2.1 
Thyasira equalis 22 9.1 
Thyasira ferruginea 3 1.2 
Thyasira sp. 1 2 0.8 
Thyasira sp. 2a 2 0.8 
Thyasira sp. 3 2 0.8 
Thyasira sp. 47 b 1 0.4 
Thyasira sp. 128 1 0.4 
Mysella verrilli 2 0.8 
Mysella sp. 2 2 0.8 
Policordia densicostata 3 2 
Policordia sp. 128 8 3.9 
Cuspidaria parva 30 12.4 
Cuspidaria jeffreysi 1 0.4 
Lyonsiella abyssicola 6 2:8 
Lyonsiella fragilis 1 0.4 
Sta. 103, 2,022 m, 39°43.6’М, 70°37.4’W 
Pristigloma nitens 1 0.02 
Deminucula atacellana 4459 79.8 
Nuculoma granulosa 6 0.1 
Nuculoma similis 2 0.04 
Neilonella salicensis 2 3.9 
Ledella sublevis 4 0.07 
Yoldiella curta 308 5.4 
Yoldiella obesa obesa 28 0.5 
Malletia johnsoni 185 3:3 
Limopsis cristata affinis 189 3.4 
Dacrydium ockelmanni 10 0.02 
Bathypecten sp. a 8 1 
Limatula subovata 25 0.5 
Thyasira brevis 1 0.02 
Thyasira equalis 1 0.02 
Thyasira tortuosa 4 0.07 
Thyasira sp. 17 Е 0.1 
Thracia nitida 1 0.02 
Thracia myopsis 1 0.02 
Policordia insoleta 2 0.04 
Cuspidaria jeffreysi 1 0.02 
Cuspidaria parva 95 Th 
Lyonsiella abyssicola 48 0.9 


(continues) 


ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Sta. 115, 2,051 т, 39°39.2’N, 70°24.5’W 
Deminucula atacellana 2133 66 
Neilonella salicensis 249 (ai 
Ledella sublevis 2 0.06 
Yoldiella curta 249 Ct 
Yoldiella obesa obesa 50 1.6 
Malletia johnsoni 125 3.9 
Limopsis cristata ago. 3 Ol 
Mytilidae sp. 115 1 0.03 
Dacrydium ockelmanni 501 0.6 
Limatula subovata 92 2.9 
Pectinidae sp. b 1 0.03 
Bathypecten eucymatus И 0.2 
Thyasira croulinensis 12 0.4 
Thyasira ferruginea 19 0.5 
Thyasira tortuosa 3 OA 
Thyasira sp. 15 62 1.9 
Mysella verrilli 24 Ой 
Abra profundorum 1 0.03 
Policordia insoleta 28 0.9 
Poromya tornata 1 0.03 
Cuspidaria parva 29 0.9 
Cuspidaria sp. 2 0.06 
Rhinoclama halimera 3 0.1 
Lyonsiella formosa 1 0.03 
Lyonsiella fragilis 3 0.1 
Lyonsiella smidti 1 0.03 
Incerte cedis sp. 1 a 21 Gay 
Incerte cedis sp. 115 1 0.03 

Sta. 210, 2,064 m, 39°43.0’N, 70°46.0’W 
Deminucula atacellana 410 39.8 
Nuculoma granulosa 6 0.6 
Nuculoma similis 1 04 
Neilonella salicensis 48 4.7 
Ledella ultima 1 0.1 
Spinula hilleri 5 0.5 
Yoldiella curta 4 0,4 
Yoldiella jeffreysi 1 0.1 
Yoldiella obesa obesa 14 1.1 
Malletia johnsoni 22 2,1 
Limopsis cristata agg. 4 0.4 
Dacrydium ockelmanni 817 9.0 
Limatula subovata фо Ib 
Limatula sp. 3 30 2.9 
Bathypecten eucymatus 716 0.9 
Propeamussium sp. 4 0.4 
Thyasira brevis y 0.1 
Thyasira carrozae 1 0.1 
Thyasira croulinensis 30 3.4 
Thyasira equalis 12 LA 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Thyasira ferruginea 81 7.9 
Thyasira obsoleta 2 0.2 
Thyasira platyssima 2 02 
Thyasira tortuosa 36 3.9 
Thyasira verrilli 1 0.1 
Thyasira sp. 15 80 7.8 
Thyasira sp. 17 3 0.3 
Thyasira sp. 210 4 0.4 
Tellinidae sp. b 1 0.1 
Verticordia sp. 1 0.1 
Laevicordia sp. 2 OZ 
Cuspidaria atlantica 1 0.1 
Cuspidaria obesa 1 0.1 
Cuspidaria parva 14 1.4 
Cuspidaria sp. 2 1 0.1 
Rhinoclama halimera 3 0:3 
Lyonsiella abyssicola 2 0.2 
Lyonsiella perplexa 2 0.2 

Sta. 131, 2.178 m, 39°38.5'N, 70°36.5 W 
Pristigloma alba 2 0.1 
Pristigloma nitens 5 003 
Deminucula atacellana 1022 70.0 
Nuculoma granulosa 1 0.07 
Neilonella salicensis 119 8.2 
Ledella sublevis 22 1:8 
Yoldiella curta 53 3.6 
Yoldiella obesa obesa 51 39 
Malletia johnsoni 39 et 
Limopsis cristata affinis 14 1.0 
Limatula subovata de 0.8 
Bathypecten eucymatus 342 03 
Thyasira croulinensis 1 0.07 
Thyasira ferruginea 31 2.1 
Thyasira sp. 15 6 0.4 
Mysella verrilli 3 0 
Verticordia quadrata 1 0.07 
Verticordia triangularis 3 0.2 
Policordia atlantica 2 0:1 
Policordia insoleta 8 0.6 
Cuspidaria barnardi 2 0.1 
Cuspidaria parva 20 1.4 
Lyonsiella abyssicola 6 0.4 
Incerte cedis sp. 1 a 2 0.1 
Incerte cedis sp. 131 1 0.07 

Sta. 119, 2.223 т 32°15.87М, 32°16.1°W 
Brevinucula verrilli 1 2.4 
Phaseolus sp. b 2 0.4 
Malletia johnsoni 23 5.0 
Bentharca asperula 0 15,3 
Dacrydium ockelmanni 245 2.0 


(continues) 


ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Dacrydium sandersi 32 7.0 
Limea lirata 189 41.3 
Cyclopecten sp. zf 1 0.2 
Thyasira inflata 26 Dur 
Thyasira succisa atlantica 1 0.2 
Thyasira verrilli 41 9.0 
Thyasıra sp.8 a 5 4.1 
Thyasira sp. 45 6 в. 
Verticordia quadrata 2 0.4 
Cuspidaria sp. 20 4.4 
Myonera paucistriata 1 0.2 
Myonera sp. 1 02 
Incerte cedis sp. 119 3 0.7 
Sta. 62, 2,496 m, 39°26.0’М, 70°33.0’W 
Deminucula atacellana 82 21.4 
Neilonella salicensis 19 3.4 
Lametila abyssorum 2 0.5 
Ledella sublevis 3 0.8 
Yoldiella fabula 4 1.0 
Yoldiella inconspicua inconspicua 25 6.5 
Yoldiella obesa obesa 6 1.6 
Malletia johnsoni 140 36:5 
Dacrydium sandersi 69 18.0 
Thyasira sp. 15 38 9.9 
Cuspidaria parva 1 0.3 
Myonera demistriata 1 0.3 
Sta. 66, 2,802 т 38°46.7’N, 70°08.8’W 
Pristigloma nitens 3 7.8 
Yoldiella ella 1 25 
Yoldiella inconspicua inconspicua 1 2:5 
Silicula filatovae 1 29 
Malletia cuneata 1 2.5 
Malletia johnsoni 3 1:9 
Thyasira croulinensis 4 10.0 
Thyasira ferruginea 2 3.0 
Mysella sp. 1 1 2.9 
Kelliella nitida 13 32.5 
Cuspidaria curta 1 2.9 
Cardiomya costellata 1 2,8 
Cardiomya knudseni 1 2:5 
Rhinoclama notabilis 1 2:5 
Tropidomya abbreviata 1 210 
Halonympha atlantica 1 2:5 
Halonympha depressa 2 5.0 
Myonera demistriata 2 0 
Sta. 76, 2,862 т, 39°38.3’N, 67°57.8’W 
Pristigloma nitens 4 155 
Deminucula atacellana 1 0.4 
Neilonella whoii 3 qna 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Ledella sublevis 10 3.9 
Yoldiella ella 1 0.4 
Yoldiella inconspicua inconspicua 53 2048 
Yoldiella jeffreysi 32 12.4 
Limopsis tenella 2 0.8 
Dacrydium sandersi 1933 51.4 
Thyasira brevis 5 1.9 
Thyasira croulinensis 2 0.8 
Thyasira equalis 1 0.4 
Thyasira ferruginea 2 0.8 
Kelliella atlantica 2 0.8 
Polycordia jeffreysi 1 0.4 
Laevicordia horrida 4 40 
Lyonsiella perplexa 3 1.2 


Sta. 72, 2,864 m, 38°16.0’N, 71°47.0’W 
Pristigloma alba 
Pristigloma nitens 
Neilonella whoii 
Lametila abyssorum 
Ledella sublevis 
Yoldiella ella 
Yoldiella fabula 
Yoldiella inconspicua inconspicua 12 
Yoldiella jeffreysi 
Yoldiella obesa obesa 1 
Malletia abyssorum 
Malletia cuneata 
Malletia johnsoni 
Dacrydium sandersi 1 
Thyasira croulinensis 
Thyasira ferruginea 
Thyasira sp. 15 
Thyasira sp. 72 
Laevicordia horrida 
Cardiomya knudseni 
Halonympha atlanta 
Incerte cedis sp. 72 


Sta. 64, 2,886 т, 38°46.0’N, 70°06.0’W 


— 


= = © = N = OWN © = © = Oo © N = O © © ON 
© 

ее © 

w 0 O1 N O1 & O1 O1 & O1 O1 


= 
AN 
Eh 0 


2.3 


ODooooono 
OOOO SIGNO ‘Go 


Pristigloma alba 11 2.2 
Deminucula atacellana 1 0.2 
Brevinucula verrilli 12 2.5 
Neilonella salicensis 2 0.4 
Prelametila clarkei 60 ler 
Ledella sublevis Fa 1,4 
Yoldiella fabula 1 0.2 
Yoldiella inconspicua inconspicua 80 15.5 
Yoldiella jeffreysi 7 1.4 
Yoldiella obesa obesa 8 1.6 
Silicula fragilis 1 0.2 
Malletia abyssorum 5 1.0 


(continues) 


ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Malletia cuneata 6 22 
Malletia johnsoni 59 10.4 
Limopsis tenella 3 0.6 
Dacrydium sandersi 173 34.2 
Axinus grandis 1 0.2 
Thyasira brevis 22 4.3 
Thyasira croulinensis 9 1.8 
Thyasira ferruginea 1 0.2 
Mysella verrilli 4 0.8 
Kelliella nitida Г. 1.4 
Thracia nitida 1 0.2 
Laevicardia horrida 2 0.4 
Cuspidaria inflata 1 0.2 
Cardiomya knudseni S 0.6 
Myonera demistriata 24 4.7 
Halonympha atlanta 4 0.8 
Incerte cedis sp. 64 1 0.2 
Sta. 112, 2,900 т, 38°50.4’N, 69°54.7’W 
Pristigloma nitens 2 33.3 
Ledella ultima 1 16.7 
Thyasira brevis 1 16.7 
Thyasira sp. 15 2 35:3 
Sta. 340, 3,356 m, 38°14.4’N, 70°20.3’W 
Deminucula atacellana 25 6.4 
Neilonella whoii 95 24.4 
Ledella sublevis 19 3.9 
Ledella ultima 14 3.6 
Yoldiella ella 16 4.1 
Yoldiella fabula 1 03 
Yoldiella inconspicua inconspicua 95 24.4 
Yoldiella jeffreysi 18 3 
Malletia abyssorum 65 107 
Malletia cuneata 43 1 
Bentharca asperula 3 0.8 
Limatula laminifera 1v 0.3 
Limatula louiseae 1v 0.3 
Poromya tornata 2 0.5 
Муопега demistriata 1 0.3 
914.99, os 031m, oo 02.0 N, 68°32.0W 
Pristigloma alba 4 19 
Tindaria callistiformis 3 0.9 
Tindaria miniscula 21 6.6 
Ledella sublevis 1 oS) 
Ledella ultima 195 60.9 
Yoldiella fabula > 0.9 
Yoldiella inconspicua inconspicua 3 0.9 
Yoldiella obesa obesa 3 0.9 
Malletia polita 59 18.4 
Hyalopecten sp. a 1 0.3 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


No. of Occurrence 


Sample Individuals in % 
Parvamussium sp. Z 4 4:3 
Thyasira brevis 12 38 
Thyasira ferruginea 2 0.6 
Thyasira sp. 15 4 0.9 
Policordia densicostata 1 0.3 
Cuspidaria sp. (shell dissolved) 2 0.6 
Myonera demistriata 1 0:3 
Incerte cedis sp. 95 1 0.3 
Sta. 77, 3,806 m, 38°00.7’N, 69°16.0’W 
Solemya grandis 6 0.2 
Deminucula atacellana 54 1.8 
Brevinucula verrilli 11 0.4 
Tindaria callistiformis 622 20.3 
Neilonella whoii 753 24.5 
Ledella ultima 1423 46.3 
Yoldiella ella 4 0.1 
Yoldiella fabula 1 0.03 
Yoldiella jeffreysi 109 3.6 


Malletia abyssorum 10 
Malletia cuneata 6 
Bathyarca inaequisculpta 1 
Limopsis galathea 11 0.4 
Limopsis tenella 56 
Poromya tornata 1 
Verticordia triangularis 4 


Sta. 126, 3,806 m, 39°37.0'N, 66%47.0'W 


Pristigloma alba 2 0.2 
Pristigloma nitens 4 0.3 
Deminucula atacellana 3 03 
Lametila abyssorum 6 9:5 
Ledella sublevis 8 0.7 
Ledella ultima 56 4.8 
Yoldiella dissimilis 8 0.7 
Yoldiella ella 4 0.3 
Yoldiella inconspicua inconspicua 14 1:2 
Yoldiella jeffreysi 138 418 
Silicula fragilis 41 38 
Malletia abyssorum 139 11.9 
Bentharca asperula 504 43.0 
Bathyarca inaequisculpta 10 0.9 
Limopsis galathea 1 0.1 
Limopsis tenella 42 3.6 
Hyalopecten undatus 61 92 
Parvamussium sp. d 13 1.1 
Thyasira ferruginea 6 0.5 
Thyasira transversa 54 4.6 
Mysella sp. 1 1 0.1 
Policordia gemma 5 0.4 
Policordia insoleta 9 0.8 
Cuspidaria parva 28 2.4 


(continues) 


100 ALLEN 


(continued) 


No. of Occurrence 


Sample Individuals in % 
Cuspidaria sp. 126 1 0.1 
Cardiomya knudseni 1 0.1 
Cardiomya sp. 1 0.1 
Myonera octoporosa 2 Gx 
Bidentaria atlantica 2 0.2 
Myonera paucistriata 1 0.1 
Lyonsiella abyssicola 5 0.4 
Lyonsiella fragilis 1 0.1 
Sta. 78, 3,828 т, 38°00.8’N, 69°18.7 W 
Pristigloma nitens 4 0.4 
Deminucula atacellana 12 12 
Brevinucula verrilli 6 0.6 
Tindaria callistiformis 18 1.8 
Neilonella whoii 199 20.4 
Ledella ultima 250 29.6 
Yoldiella ella 11 1.1 
Yoldiella fabula 6 0.6 
Yoldiella inconspicua inconspicua € 078 
Yoldiella jeffreysi 62 6.3 
Silicula filatovae 10 1.0 
Malletia abyssorum 144 14.7 
Malletia cuneata 64 6.8 
Limopsis galathea 1 0 
Limopsis tenella 15 1.5 
Thyasira brevis 3 0.3 
Kelliella nitida 1 0.1 
Verticordia triangularis 5 0.5 
Policordia gemma 1 0.1 


Sta. 85, 3,834 m, 37°59.2’М, 69°26.2W 


Pristigloma alba 4 0.06 
Pristigloma nitens 4 0.06 
Deminucula atacellana 69 1.1 
Brevinucula verrilli 21 0.3 
Tindaria callistiformis 882 138 
Neilonella whoii 1150 18.0 
Pseudotindaria erebus 1 0.02 
Lametila abyssorum 60 0.9 
Ledella aberrenta 1 0.02 
Ledella sublevis 3 0.05 
Ledella ultima 1045 16.4 
Yoldiella americana 6 0.09 
Yoldiella ella 12 0.2 
Yoldiella fabula 18 0.2 
Yoldiella inconspicua inconspicua 32 0.5 
Yoldiella jeffreysi 413 6.5 
Silicula filatovae 55 0.9 
Silicula fragilis 39 0.6 
Malletia abyssorum 1475 291 
Malletia cuneata 789 12:4 
Malletia johnsoni 2 0.03 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Malletia pallida 
Bathyarca inaequisculpta 
Limopsis galathea 
Limopsis tenella 
Dacrydium abyssorum 
Limatula margaretae 
Parvamussium sp. Z 
Thyasira brevis 
Thyasira transversa 
Thyasira sp. 17 
Montacuta sp. 1 
Kelliella nitida 
Verticordia quadrata 
Verticordia triangularis 
Policordia gemma 
Myonera angularis 
Myonera octoporosa 
Lyonsiella smidti 


Sta. 175, 4,693 т, 36°36.0’М, 68°29.0’W 
Pristigloma alba 
Pristigloma nitens 
Tindaria callistiformis 
Ledella ultima 
Yoldiella americana 
Yoldiella jeffreysi 
Malletia abyssorum 
Malletia cuneata 
Malletia pallida 
Limopsis galathea 
Hyalopecten sp. a 
Thyasira ferruginea 


Sta. 70, 4,680 m, 36°23.0’М, 67°58.0’W 
Pristigloma nitens 
Ledella sublevis 
Ledella ultima 
Yoldiella americana 
Yoldiella fabula 
Malletia abyssorum 
Malletia cuneata 
Bentharca asperula 
Limopsis galathea 
Dacrydium abyssorum 
Thyasira brevis 
Thyasira sp. 40 
Kelliella nitida 
Cuspidaria sp. 
Myonera atlantica 
Myonera octoporosa 


No. of 
Individuals 


Occurrence 
in % 


(continues) 


101 


102 ALLEN 


(continued) 


Sample 


Sta. 92, 4,694 m, 36°20.0’N, 67°56.0’W 
Ledella ultima 
Yoldiella americana 
Yoldiella fabula 
Yoldiella subcircularis 
Malletia abyssorum 
Malletia cuneata 
Bentharca asperula 
Limopsis galathea 
Hyalopecten sp. a 
Myonera octoporosa 
Incerte cedis sp. 1 a 


Sta. 108, 4,739 m, 36°24.0’N, 68°04.8’W 
Ledella ultima 
Limopsis galathea 


Sta. 101, 4,740 m, 36°24.2’N, 68°01.3'W 
Ledella ultima 
Yoldiella americana 


Sta. 100, 4,743 т, 33%56.8'N, 65°47.0 W 
Yoldiella americana 
Yoldiella inconspicua inconspicua 
Yoldiella subcircularis 
Limatula louiseae 
Hyalopecten sp. a 
Astarte sp. 
Incerte cedis sp. 100 


Sta. 109, 4,750 m, 36°25.0’N, 68°06.0’W 
Pristigloma nitens 
Ledella ultima 
Yoldiella americana 
Malletia abyssorum 
Malletia polita 
Mysella sp. 1 


Sta. 84, 4,794 m, 36°24.4’N, 67°56.0’W 
Pristigloma nitens 
Brevinucula verrilli 
Tindaria callistiformis 
Tindaria perrieri 
Ledella ultima 
Yoldiella americana 
Yoldiella fabula 
Yoldiella jeffreysi 
Yoldiella subcircularis 
Silicula filatovae 
Malletia abyssorum 
Malletia cuneata 
Malletia pallida 
Bentharca asperula 
Limopsis galathea 


No. of 
Individuals 


209 


Occurrence 
in % 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Dacrydium abyssorum 
Thyasira robusta 
Thyasira transversa 
Thyasira sp. 9 
Kelliella nitida 
Cuspidaria undata 
Cuspidaria sp. 
Myonera octoporosa 
Myonera sp. 
Edentaria simplis 


Sta. 121, 4,800 m, 35°50.0’М, 65°11.0°W 
Pristigloma alba 
Pristigloma nitens 
Tindaria callistiformis 
Ledella ultima 
Ledella sp. 

Spinula sp. 

Yoldiella americana 
Malletia abyssorum 
Malletia pallida 
Thyasira transversa 
Epilepton elpis 
Myonera octoporosa 
Incerte cedis sp. 121 


Sta. 125, 4,825 m, 37°24.0’М, 65°54.0’W 
Pristigloma alba 
Ledella ultima 
Yoldiella americana 
Yoldiella fabula 
Silicula filatovae 
Malletia abyssorum 
Malletia pallida 
Limopsis galathea 
Thyasira ferruginea 
Thyasira transversa 
Abra profundorum 
Cuspidaria sp. 
Myonera octoporosa 
Edentaria simplis 


Sta. 122, 4,833 m, 35°50.0’N, 64°57.5’W 
Pristigloma alba 
Pristigloma nitens 
Tindaria callistiformis 
Ledella ultima 
Yoldiella americana 
Yoldiella subcircularis 
Malletia abyssorum 
Malletia pallida 
Thyasira transversa 
Epilepton elpis 


No. of 
Individuals 


Gores 
ыы AN © © N 9 


O) 
N 
== ND © © 


O) 
©) 


178 


— 
+ 


Occurrence 
in % 


DO00N00N0S008WO 
NOUS ANDO On 


(continues) 


103 


104 


ALLEN 


(continued) 


Sample 


Myonera octoporosa 
Incerte cedis sp. 121 
Incerte cedis sp. 122 


Sta. 123, 4,853 т, 37°29.0'N, 64°14.0 W 
Pristigloma nitens 
Tindaria callistiformis 
Ledella ultima. 
Ledella sp. 
Spinula sp. 
Yoldiella americana 
Yoldiella jeffreysi 
Malletia abyssorum 
Thyasira brevis 


Sta. 124, 4,862 m, 37°26.0'N, 63°59.5’W 
Tindaria callistiformis 
Neilonella whoii 
Ledella ultima 
Yoldiella americana 
Yoldiella jeffreysi 
Malletia abyssorum 
Malletia pallida 
Thyasira atlantica 
Thyasira transversa 
Abra profundorum 


Sta. 80, 4,970 m, 34°49.8’N, 66°34.0’W 
Neilonella whoii 
Ledella ultima 
Yoldiella americana 
Yoldiella subcircularis 
Silicula filatovae 
Malletia abyssorum 
Malletia pallida 
Bentharca asperula 
Dacrydium abyssorum 
Abra profundorum 


Sta. 98, 4,977 m, 34°45.5’N, 66°30.5’W 
Yoldiella subcircularis 
Malletia polita 
Thyasira sp. 98 
Verticordia sp. 98 


Sta. 99, 4,977 т, 34°43.0’М, 66°23.3’W 
Tindaria callistiformis 
Yoldiella subcircularis 
Silicula filatovae 
Malletia polita 
Sta. 83, 5,000 т, 34°46.5’N, 66°30.0’W 
Ledella sp. 
Spinula sp. 
Yoldiella americana 


No. of 
Individuals 


Occurrence 
in % 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Yoldiella similirus 
Silicula mcalesteri 
Malletia abyssorum 
Malletia pallida 
Limopsis galathea 
Dacrydium abyssorum 
Abra profundorum 
Cuspidaria sp. 
Myonera octoporosa 
Lyonsiella smidti 


Sta. 93, 5,007 m, 34°39.0’М, 66°26.0’W 
Ledella ultima 
Yoldiella americana 
Yoldiella subcircularis 
Silicula filatovae 
Malletia abyssorum 
Malletia pallida 
Bentharca asperula 
Dacrydium abyssorum 
Lyonsiella smidti 


Sta. 120, 5,023 т, 34°43.0'N, 66°32.8" W 
Cuspidaria atlantica 


Sta. 81, 5,042 т, 34°41.0’N, 66°28.0’W 
Tindaria callistiformis 
Ledella ultima 
Yoldiella americana 
Yoldiella similirus 
Silicula mcalesteri 
Malletia abyssorum 
Limopsis galathea 


SURINAM BASIN 


Sta. 297, 523 m, 07°45.3’М, 54*24.0'W 
Solemya sp. 293 
Pristigloma sp. a 
Nuculoma elongata 
Neilonella salicensis 
Malletia grasslei 
Malletia malita 
Malletia surinamensis 
Limopsis surinamensis 
Limatula celtica 
Propeamussium Sp. C 
Cyclopecten simplex 
Cyclopecten sp. c 
Myrtea lens 
Thyasira croulinensis 
Thyasira ferruginea 
Malletia succisa atlantica 
Thyasira transversa 


No. of 
Individuals 


NO 
2 N © ND NN © © © N NY 


ES 
MPMOANAOW — 


Occurrence 
in % 


NO 


> — 
a — — (© ©) = N 9 — 
© © © O1 O1 © © © © 


(continues) 


105 


106 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Thyasira sp. 15 12 0.6 
Thyasira sp. 297 1 0.05 
Abra profundorum 4 0.2 
Tellinidae sp. a 1 0.05 
Kelliella atlantica 820 38.4 
Kelliella elongata 1 0.05 
Thracia durouchouxi 1 0.05 
Policordia sp. 297 4 0.2 
Cuspidaria parva 149 7.0 
Cuspidaria sp. p 5 0.2 
Cuspidaria sp. q 3 0.1 
Cardiomya costellata 17 0.8 
Myonera paucistriata 6 0.3 
Sta. 295, 1,022 m, 08°04.2’N, 54°21.3’W 
Solemya sp. 1 0.04 
Pristigloma sp. a 58 2.1 
Microgloma sp. $ 189 6.9 
Nuculoidea bushae 28 150 
Neilonella salicensis 1575 DS 
Ledella acinula 55 23 
Yoldiella biguttata 2 0.07 
Yoldiella curta 25 0.9 
Malletia grasslei ie 9.5 
Malletia таШа 20 ony 
Malletia surinamensis 40 1.5 
Limopsis cristata intermedia 37 1.4 
Parvamussium lucidum 1 0.04 
Propeamussium sp. C 3 0.1 
Cyclopecten pustulosus 32 12 
Thyasira croulinensis 28 10 
Thyasira equalis о 0.1 
Thyasira ferruginea 30 ard 
Thyasira succisa atlantica Fa 2.6 
Thyasira sp. 2 с 5 0.2 
Thyasira sp. 4 2 0.07 
Thyasira sp. 17 1 0.04 
Tellinidae sp. a 1 0.04 
Kelliella atlantica 245 8.9 
Poromya sp. 1 0.04 
Cuspidaria atlantica 5 0.2 
Cuspidaria parva 1 0.04 
Myonera paucistriata 254 9.2 
Cuspidaria sp. 24 0.9 


Sta. 293, 1,518 m, 08°58.0’N, 54°04.3’W 
Solemya sp. 293 2 
Pristigloma sp. a 4 
Microgloma sp. $ 151 
Deminucula atacellana 9 
Nuculoidea pernambucensis 23 
Neilonella salicensis 2 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Tindariopsis agatheda 
Ledella acinula 
Ledella jamesi 
Yoldiella biguttata 
Yoldiella curta 
Yoldiella fabula 
Yoldiella perplexa 
Portlandia sp. 

Malletia malita 
Limopsis cristata intermedia 
Limatula margaretae 
Propeamussium sp. а 
Kelliella adamsi 
Kelliella atlantica 


Sta. 299, 2,076 m, 07°55.1'N, 55°42.0’W 
Microgloma yongei 
Microgloma sp. s 
Deminucula atacellana 
Brevinucula verrilli 
Neilonella salicensis 
Tindariopsis aeolata 
Tindariopsis agatheda 
Tindariopsis sp. 

Ledella acinula 
Ledella jamesi 
Ledella solidula 
Yoldiella biguttata 
Yoldiella perplexa 
Yoldiella sinuosa 
Portlandia sp. 
Limatula louiseae 
Bathypecten sp. a 
Limopsis cristata intermedia 
Thyasira ferruginea 
Thyasira succisa atlantica 
Thyasira transversa 
Thyasira verrilli 
Thyasira sp. 4 
Thyasira sp. 17 
Thyasira sp. 30 
Thyasira sp. 299 
Kelliella atlantica 
Abra profundorum 
Abra sp. 

Cuspidaria sp. L 
Incerte cedis sp. 299 

Sta. 301, 2,500 m, 08°12.4’N, 55°50.2’W 
Microgloma yongei 
Microgloma sp. $ 
Deminucula atacellana 


No. of 
Individuals 


14 
4 
2 

18 

49 
6 

22 
4 
2 

44 


© 


— 
а ор 


136 
48 
12 


Occurrence 
in % 


3:9 
VE 
0.6 


(continues) 


107 


108 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Brevinucula verrilli 18 1.8 
Neilonella whoii 29 253 
Tindariopsis agatheda 7 0.7 
Tindariopsis aeoleta 5 05 
Ledella jamesi 51 5,2 
Spinula subexisa 2 02 
Yoldiella biguttata 44 4.4 
Yoldiella enata 178 17.7 
Yoldiella ovata 324 327 
Bathyarca inaequisculpta 20 2:0) 
Parvamussium obliquum 8 0.8 
Thyasira ferruginea 1 0.1 
Thyasira inflata 25 2.8 
Thyasira succisa atlantica 13 128 
Thyasira verrilli 15 1.5 
Thyasira sp. 45 1 0.4 
Epilepton sp. 21 2 0.2 
Mysella verrilli 2 02 
Abra profundorum oF ЭХ 
Kelliella abyssicola 19 1:9 
Poromya sp. 301 1 0.1 
Incerte cedis sp. 301 3 0.3 
Sta. 303, 2,853 m, 08°28.8’N, 56°04.5 W 
Solemya sp. 1 0.1 
Microgloma turnerae 1 0.1 
Microgloma yongei 163 24.0 
Microgloma sp. s 5 07 
Deminucula atacellana T 1.0 
Nuculoma granulosa 1 0.1 
Brevinucula verrilli 9 1.3 
Tindaria miniscula 1 0.1 
Neilonella whoii 8 e 
Prelametila clarkei 2 0:8 
Ledella jamesi 268 69.5 
Yoldiella biguttata 4 0.6 
Yoldiella ella 12 1.8 
Yoldiella enata 23 3.4 
Yoldiella fabula 2 0:3 
Yoldiella frigida 1 0.1 
Yoldiella jeffreysi fe 140 
Yoldiella ovata 22 eee 
Bathyarca inaequisculpta 58 8.6 
Limopsis surinamensis 5 0.7 
Limatula margaretae 3 0.4 
Parvamussium lucidum 1 0.1 
Propeamussium sp. с 1 0.1 
Thyasira transversa 2 0.3 
Abra profundorum 2 0.3 
Kelliella abyssicola о Di 
Kelliella atlantica 64 9.4 
Cuspidaria obesa 1 0.1 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 109 


(continued) 


No. of Occurrence 
Sample Individuals in % 


Sta. 306 3,429 т, 09°31.1’N, 56°20.6 W 


Pristigloma nitens 2 0.9 
Brevinucula verrilli 27 ire 
Neilonella whoii 1 0.4 
Spinula hilleri 5 2:2 
Yoldiella jeffreysi 38 16:7 
Bathyarca inaequisculpta 41 18.0 
Dacrydium abyssorum 2 0.9 
Propeamussium Sp. с 4 1.8 
Thyasira biscayensis 1 0.4 
Thyasira inflata 93 40.8 
Thyasira sp. 306 1 0.4 
Abra profundorum 1 0.4 
Kelliella abyssicola 11 4.8 
Verticordia quadrata 1 0.4 
Laevicordia horrida 2 0.9 
Myonera atlantica 3 133 
Sta. 307, 3,835 т, 12°35.4’N, 58°59.3’W 
Solemya sp. 1 0.6 
Tindaria callistiformis 1 0.6 
Neilonella salicensis 2 ng 
Neilonella whoii 15 9.2 
Pseudotindaria erebus 22 13.4 
Ledella ultima 6 on 
Spinula scheltemae 10 6.1 
Yoldiella jeffreysi 30 18.3 
Malletia polita 1 0.6 
Bathyarca inaequisculpta 48 29.3 
Limopsis galathea 26 15:89 
Kelliella atlantica 2 2 
Sta. 291, 3,868 m, 10%06.1'N, 55°14.0’W 
Brevinucula verrilli 10 1.9 
Neilonella whoii 43 8.2 
Yoldiella fabula 1 0.2 
Yoldiella jeffreysi 158 29.4 
Portlandia sp. 3 0.6 
Bentharca asperula 1 0.2 
Bathyarca inaequisculpta 1 0.2 
Bathyarca pectunculoides 1 0.2 
Limopsis surinamensis 5 1.0 
Dacrydium hedleyi 24 4.6 
Halopecten undatus 1 0.2 
Parvamussium sp. q 2 0.4 
Bathypecten eucymatus 6 142 
Thyasira pygmaea 1 0.2 
Thyasira tortuosa 1 0.2 
Thyasira transversa 54 103 
Thyasira sp. 1 11 2 
Thyasira sp. 15 1 0.2 
Abra profundorum 30 57 


(continues) 


ALLEN 


(continued) 


No. of Occurrence 


Sample Individuals in % 
Kelliella abyssicola 4 0.8 
Verticordia quadrata 5 1.0 
Verticordia sp. 2 0.4 
Laevicordia horrida 7 1.3 
Роготуа granulata 1 0.2 
Cuspidaria sp. а 4 0.8 
Cuspidaria sp. 1 131 24.9 
Cuspidaria sp. 2 4 0.8 
Tropidomya abbreviata 8 eo 
Lyonsiella smidti 2 0.4 
Incerte cedis sp. 291 a 3 0.6 
Incerte cedis sp. 291 b 5 1.0 
Sta. 288, 4,429 m, 11°02.2’N, 55°05.5’W 
Tindaria callistiformis 13 2.4 
Neilonella whoii 19 3.0 
Pseudotindaria erebus 9 ПК 
Ledella ultima 51 9.6 
Yoldiella americana 45 8.4 
Yoldiella jeffreysi 91 5.0 
Yoldiella subcircularis 4 8.3 
Malletia polita 71 198 
Bathyarca inaequisculpta 44 8.3 
Limopsis galathea 111 20.8 
Dacrydium abyssorum 14 246 
Dacrydium hedleyi 9 ik 
Propeamussium thalassinum 1 0.2 
Thyasira tortuosa 4 0.8 
Thyasira sp. 1 1 0.2 
Thyasira sp. 2 a 4 0.8 
Thyasira sp. 3 1 0.2 
Kelliella atlantica 61 11.4 
Sta. 287, 4,980 т 13°16.0’N, 54°52.2’W 
Tindaria callistiformis 10 07 
Neilonella whoii 2 5 
Pseudotindaria erebus 87 6.2 
Ledella ultima 84 6.0 
Spinula scheltemae 1 0.1 
Yoldiella americana 12 0.9 
Yoldiella jeffreysi 5 0.4 
Yoldiella subcircularis 5 0.4 
Malletia abyssorum 5 0.4 
Malletia polita 180 12.8 
Bathyarca inaequisculpta 108 7-7: 
Limopsis galathea 621 44.0 
Dacrydium abyssorum 2% 1.9 
Dacrydium hedleyi 2 0.1 
Cyclopecten pustulosus 3 0.2 
Cyclopecten sp. ze 1 0.1 
Thyasira transversa 14 170 
Abra profundorum 30 an 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Kelliella atlantica 
Cuspidaria sp. 
Cardiomya sp. 
Myonera atlantica 
Myonera sp. 


Sta. Biovema DS01, 5,097 m 10°57.2’N, 
45°07.6W 

Pristigloma alba 

Ledella ultima 

Malletia polita 


Sta. Biovema DS04, 5,100 m 10°46.3'N, 

42°41.4’W 
Pristigloma alba 
Tindaria callistiformis 
Neilonella whoii 
Ledella ultima 
Silicula filatovae 
Yoldiella americana 
Malletia abyssorum 
Malletia polita 
Limopsis cristata agg. 
Dacrydium sp. 
Limatula louiseae 
Thyasira sp. 1 
Thyasira sp. 2 
Cuspidaria sp. 
Incerte cedis 

Sta. Biovema DS05, 5,100 m, 10%46.0'N, 

42°40.3"W 
Pristigloma nitens 
Neilonella whoii 
Ledella ultima 
Silicula filatovae 
Yoldiella americana 
Yoldiella subcircularis 
Malletia polita 
Bathyarca inaequisculpta 
Limatula louiseae 
Thyasira transversa 
Mysella sp. 
Kelliella elongata 


Sta. Biovema DSO3, 5,150 m, 10%47.1'N, 
42°40.7'W 

Neilonella hampsoni 

Ledella ultima 

Yoldiella americana 

Yoldiella subcircularis 

Malletia abyssorum 

Malletia polita 


No. of 
Individuals 


135 
2 


1 
1 
5 


№ > = 


oF — 


> 
== —= O1 O1 N NN = © O1 ND 0 == 01 


— 


Occurrence 
in % 


9.6 
0.1 
0.1 
0.1 
0.4 


(continues) 


111 


eZ 


ALLEN 


(continued) 


Sample 


CAPE VERDE BASIN 


Sta. Biovema DS11, 5,867 т, 11°37.5’N, 
32/5368 VV 

Ledella ultima 

Spinula hilleri 

Yoldiella curta 

Malletia polita 

Thyasıra transversa 

Kelliella atlantica 


Sta. Biovema 0309, 5,875 m, 11°36.4°N, 
32°51,8 W 

Ledella ultima 

Spinula sp. 

Yoldiella curta 

Malletia pallida 

Limopsis sp. 

Thyasira subovata 

Kelliella sp. 


Sta. Biovema DS10, 5,875 m, 11°36.8’М, 
32 °52.5’W 

Ledella ultima 

BRAZIL BASIN 


Sta. 168, 416 m, 07°50.0'S, 34°28.0 W 
Propeamussium Sp. 
Thracia Sp. 
Myonera Sp. 


Sta. 169, 827 m, 08°03.0'S, 34°23.0W 
Nuculoidea pernambucensis 
Yoldiella curta 
Dacrydium sandersi 
Protocuspidaria verityi 
Lyonsiella subquadrata 


Sta. 167, 1,007 т, 07°58.0’S, 34°17 W 
Microgloma sp. s 
Nuculoidea pernambucensis 
Brevinucula subtrangularis 
Tindariopsis agatheda 
Ledella acinula 
Ledella oxira 
Yoldiella biguttata 
Yoldiella curta 
Dacrydium sandersi 
Parvamussium lucidum 
Parvamussium sp. a 
Bathypecten sp. a 
Axinodon symmetros 
Policordia laevis 
Protocuspidaria verityi 


No. of 
Individuals 


N © 
© = = ND = © © À O1 NN = = = À O1 


Occurrence 
in % 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 
Cuspidaria parva 
Cuspidaria sp. 
Myonera sp. 


Lyonsiella subquadrata 


Sta. 162, 1,493 т, 08°02.0’S, 34°03.0W 
Neilonella corpulenta 
Ledella oxira 
Parvamussium lucidum 
Cuspidaria circinata 
Myonera sp. 


Sta. 156, 3,495 т, 00°46.0’S, 29°28.0’W 
Pristigloma alba 
Pristigloma nitens 
Nuculoma callicredemna 
Pseudotindaria erebus 
Lametila abyssorum 
Ledella ultima 
Spinula hilleri 
Yoldiella ella 
Bentharca asperula 
Limopsis tenella 
Dacrydium sandersi 
Limatula louiseae 
Hyalopecten sp. e 
Parvamussium sp. a 
Bathypecten sp. a 
Cyclopecten sp. zg 
Thyasira transversa 
Thyasira sp. 8 a 
Cuspidaria parva 
Cuspidaria sp. 
Rhinoclama notabilis 
Myonera octoporosa 
Myonera sp. 1 
Myonera sp. 2 
Poromya tornata 
Verticordia quadrata 
Policordia gemma 
Poromya sp. 
Lyonsiella formosa 


Sta. 155, 3,783 m, 00°03.0’S, 27°48.0’W 
Brevinucula verrilli 
Neilonella hampsoni 
Pseudotindaria erebus 
Lametila abyssorum 
Spinula hilleri 
Yoldiella ella 
Bentharca asperula 
Bathyarca inaequisculpta 


No. of 
Individuals 


4 
1 


4 


Oo — № = — 


© 


SS 
BAA = = OHH = WNANONNFHDAPHWOHANNANN 


—Dh 


NY & © © © N 


Occurrence 
in % 


(continues) 


118 


114 ALLEN 


(continued) 


No. of 
Sample Individuals 


Limopsis galathea 27. 
Limopsis tenella 24 
Dacrydium sandersi 90 
Limatula celtica 

Limatula louiseae 

Pectinidae sp. c 

Pectinidae sp. d 

Pectinidae sp. 

Hyalopecten sp. e 

Thyasira sp. 17 

Verticordia quadrata 

Myonera sp. 1 1 
Myonera sp. 2 

Myonera sp. 3 

Poromya tornata 

Cetoconcha braziliensis 

Lyonsiella formosa 


ARGENTINE BASIN 
Sta. 284, 98 m, 36°08.3’S, 53°42.3°W 


EN SS м“ ED 


Cardiidae sp. 4 6 
Sta. 282, 165 m, 3617.95, 53°31.3°W 
Zygochlamys patagonia 3 
Sta. 280, 293 m, 36°17.0°S, 53°23.9W 
Solemya sp. 280 1 
Nuculidae sp. 280 2 
Nuculoma perforata 15 
Yoldiella robusta 3495 
Silicula mcalesteri 39 
Lucinidae sp. 280 1 
Thyasira croulinensis 15 
Thyasira equalis 31 
Thyasira ferruginea 60 
Thyasira sp. 280 18 
Thyasira sp. 2 a 238 
Aesthenotherus hempilli йе 
Veneridae sp. 280 1 
Sia. 236, 910 m, oo 27.195, 53 31.0 W 
Solemya sp. 237 1 
Thyasira sp. 346 c 2 
Carditidae sp. 5 36 
Leptonidae sp. v 2 
Kelliella sp. 236 1 
Incerte cedis sp. 236 1 
Incerte cedis sp. 237 2 


sa.237..1.041m.30°32.65, 03 230 W 


Solemya sp. 237 1 
Nuculoma perforata $ 
Propoleda carpenteri 196 


Occurrence 
in % 


13.8 
124 
45.2 

les) 

ESS) 
0.05 
0.05 
0.05 
0.05 

A) 
0.05 

7.0 
0.05 
0.05 

1.0 
0.05 
0.05 


100.0 
100.0 


0.03 
0.05 
0.4 
89.1 
и 
0.03 
0.4 
0.8 
ANS 
0.5 
6.1 
0.2 
0.03 


2.2 
4.4 
80.0 
4.4 
2 
22 
4.4 


0.2 
12 
32:2 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Yoldiella argentinea 26 4.3 
Yoldiella similirus > 8.0 
Limea argentineae 2 0.3 
Thyasira croulinensis Э 0.5 
Thyasira equalis 1 0.2 
Thyasira transversa 5 0.5 
Neolepton profundorum 173 28.5 
Carditidae sp. 1 64 10.5 
Cuspidaria sp. 1 7) 0.3 
Cuspidaria sp. 2 32 Did 
Rhinoclama notabilis 127 20.9 
Incerte cedis sp. 237 1 0.2 

Sta. 239, 1,679 т, 36°49.0'S, 53°15.4W 
Solemya sp. 259 2 (OS 
Nuculoma perforata 37 1.8 
Ledella jamesi 889 44.6 
Ledella sublevis 142 re 
Propoleda carpenteri 8 0.4 
Yoldiella argentinea 26 13 
Yoldiella curta 36 1.8 
Yoldiella sp. 6 0.3 
Limopsis sp. 239 2 0.1 
Dacrydium ockelmanni 9 0:5 
Dacrydium sp. L 1 0.05 
Limea argentineae 6 0.3 
Bathypecten sp. d 268 ta 
Thyasira alleni 1 0.05 
Thyasira carrozae 196 9.7 
Thyasira croulinensis 13 0.7 
Thyasira ferruginea 19 0.9 
Thyasira transversa 169 8.4 
Thyasira sp. 239 2 (00 
Mysella verrilli 76 3.8 
Kelliella elongata 13 07 
Cardiomya knudseni 1 0.05 
Tropidomya abbreviata 90 4.5 
Lyonsiella sp. 239 1 0.05 

Sta. 264, 2,048 m 36°12.7'S, 52°42.7’W 
Nuculoma perforata 1 0.4 
Ledella jamesi 8 3.0 
Ledella sublevis 2 ВЕ 
Yoldiella curta 10 35 
Yoldiella inconspicua profundorum 6 2,2 
Limopsis spicata 21 7.9 
Thyasira carrozae 20 7.0 
Thyasira equalis 1 0.4 
Thyasira ferruginea 10 38 
Thyasira succisa atlantica 5 1.9 
Thyasira transversa 169 0923 
Thyasira sp. 15 2 0.7 


(continues) 


ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals In % 

Thyasira sp. 30 12 4.5 

Sta. 240, 2,323 m, 36°53.4’S, 53°10.2W 
Solemya sp. 280 1 05 
Nuculoma perforata 11 2.8 
Ledella jamesi 61 199 
Ledella sublevis 18 4.6 
Propoleda carpenteri 3 0.8 
Yoldiella argentinea 1 0.8 
Yoldiella curta 1 0.3 
Yoldiella robusta 2 0.8 
Limopsis cristata agg. 6 ide: 
Dacrydium ockelmanni 2 0.5 
Limea sp. 240 1 0.3 
Cyclopecten sp. a 70 176 
Thyasira carrozae 6 16 
Thyasira croulinensis 58 14.8 
Thyasira eumyaria 1 0.3 
Thyasira ferruginea 13 38 
Thyasira hydroida 3 0.8 
Thyasira obsoleta 2 0.5 
Thyasira pygmaea 84 21.4 
Thyasira subequatoria 3 0.8 
Thyasira subovata atlantica 1 0.3 
Thyasira transversa 5 Wes 
Thyasira trisinuta 1 0.3 
Thyasira sp. 15. 1 0.3 
Thyasira sp. 45. 1 0 
Thyasira sp. 47 а 2 0.5 
Neolepton profundorum 8 2.0 
Kelliella elongata 1 03 
Kelliella sp. 245 3 0.8 
Verticordia sp. 240 i 0:3 
Cuspidaria sp. 239 3 0.8 
Cardiomya gemma 1 0.3 
Cardiomya knudseni 2 0.5 
Tropidomya abbreviata 1 0.3 
Myonera atlantica 2 0:5 
Myonera paucistriata 1 0.3 
Incerte cedis sp. 240 5 1.3 
Incerte cedis 1 0.3 

Sta. 262, 2,480 т, 36°05.2’S, 52°17.9’W 
Solemya sp. 259 2 0.2 
Nuculoma perforata 1 0.1 
Phaseolus sp. с 2 0.2 
Ledella jamesi 6 0.7 
Ledella sublevis 220 24.6 
Malletia cuneata 234 23,8 
Limopsis spicata 11 eZ 
Bathypecten sp. a 6 0.7 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Propeamussium sp. C i 0.8 
Thyasira brevis 267 29.8 
Thyasira ferruginea de 0.8 
Thyasira subovata subovata 1 0.1 
Thyasira succisa atlantica 20 Zz 
Thyasira transversa 02 5.8 
Thyasira sp. 30 21 2.3 
Kelliella atlantica 1 0.1 
Thracia nitida 3 0:3 
Protocuspidaria verityi 2 0.2 
Cardiomya knudseni 7 0.8 
Myonera atlantica 1 0.1 
Myonera sp. 1 7 1.9 
Myonera sp. 2 7 0.8 
Incerte cedis sp. 262 1 0.1 

Sta. 245, 2,707 m, 36°55.7’S, 53°01.4W 
Solemya sp. 259 2 Ors 
Deminucula atacellana 17 10.2 
Ledella sublevis 8 41 
Yoldiella biguttata 2 0.3 
Yoldiella blanda 1 0.1 
Yoldiella extensa 245 32.4 
Yoldiella inconspicua profundorum 2 0.3 
Malletia cuneata 62 8.2 
Bathyarca sp. 245 2 03 
Dacrydium ockelmanni 21 2:0 
Limatula subovata 98 1 
Bathypecten sp. a 12 1:6 
Thyasira brevis 74 9.8 
Thyasira carrozae 4 08 
Thyasira equalis 1 an 
Thyasira sp. 15 1 q 
Thyasira sp. 17 1 0.1 
Neolepton profundorum tal WES 
Kelliella atlantica 10 1:6 
Kelliella sp. 245 1 0.1 
Cuspidaria barnardi 1 0.1 
Cuspidaria circinata 1 0.1 

Sta. 259, 3,977.1; 37 13.35, 5245. 0W 
Solemya sp. 259 1 0.07 
Deminucula atacellana 862 Вл 
Tindaria callistiformis 5 0.4 
Neilonella whoii ql 0.8 


Pseudotindaria championi 5 
Ledella pustulosa argentinea 6 
Ledella sublevis 24 17 
Ledella ultima 4 
Spinula scheltemae 5 
Yoldiella blanda 20 


(continues) 


ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Yoldiella inconspicua profundorum 5 0.4 
Yoldiella jeffreysi 30 2] 
Silicula fragilis 5 05 
Malletia cuneata 106 1-5 
Bathypecten sp. a 6 0.4 
Propeamussium sp. с 10 Quí 
Cyclopecten pustulosus 9 0.6 
Thyasira brevis 29 1.8 
Thyasira equalis 3 0.2 
Thyasira ferruginea 1:3 0.9 
Thyasira (robusta) 20 1.4 
Thyasira platyssima 7 035 
Thyasira subovata subovata 96 6.8 
Kelliella atlantica Tae 8.3 
Kelliella elongata 8 0.6 
Policordia gemma 3 iz 
Cuspidaria parva 1 0.07 
Cuspidaria sp. 1 0.07 
Myonera paucistriata 1 0.07 


Sta. 246, 3,343 т 37°15.1’S, 52°45.0'W 


Deminucula atacellana 16 8.4 
Tindaria miniscula 4 2.1 
Pseudotindaria erebus 1 0.5 
Ledella pustulosa argentinae 1 05 
Propeleda louiseae 3 1.6 
Yoldiella blanda 3 1.6 
Yoldiella ella 8 4.2 
Yoldiella jeffreysi 5 2.6 
Yoldiella similirus 5 2.6 
Yoldiella subcircularis 2 Я 
Yoldiella sp. 246 1 5.8 
Silicula fragilis 3 1.6 
Malletia cuneata 27 14.1 
Bathypecten eucymatus 8 4.2 
Thyasira brevis 1 0.5 
Thyasira ferruginea 4 21 
Thyasira inflata 30 157 
Thyasira subcircularis 1 0.5 
Neolepton sp. 246 2 hit 
Kelliella atlantica 30 197 
Kelliella elongata 1 0.5 
Cuspidaria sp. 2 1.4 
Cardiomya sp. 1 0,5 
Tropidomya abbreviata 3 1.6 
Lyonsiella abyssicola 8 4.2 
Sta. 243, 3,822 m, 37°36.8'$, 52°23.6’W 
Deminucula atacellana 213 67.6 
Neilonella whoii 3 1 
Ledella pustulosa argentinea 41 18 
Propeleda louiseae 2 0.6 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Yoldiella fabula 10 3.2 
Yoldiella inconspicua profundorum 1 3 
Cyclopecten sp. a 4 18 
Thyasira brevis 9 2:9 
Thyasira ferruginea 18 OF 
Thyasira robusta 1 O23 
Thyaasira succisa atlantica 1 0.3 
Thyasira transversa 2 0.6 
Thyasira sp. 243 8 2.6 
Kelliella elongata 2 0.6 

Sta. 256, 3,916 m, 37°40.9'S, 52°19.3’W 
Pristigloma nitens 3 0.2 
Deminucula atacellana 356 24.2 
Tindaria callistiformis Se 2 
Neilonella whoii 8 0.5 
Prelametila sp. 247 1 0.07 
Ledella jamesi 1 0.07 
Ledella pustulosa argentinea 97 6.6 
Ledella sublevis 3 0.2 
Ledella ultima 79 5.1 
Spinula scheltemae 237 16.1 
Propeleda louiseae 3 0 
Yoldiella blanda 63 4.3 
Yoldiella fabula 4 0.3 
Yoldiella inconspicua profundorum 19 4.1 
Malletia cuneata 350 23:8 
Malletia abyssorum 3 0.2 
Pectinidae sp. b 10 0.7 
Bathypecten eucymatus 10 0.7 
Thyasira croulinensis 36 25 
Thyasira equalis 13 0.8 
Thyasira ferruginea 42 2.9 
Thyasira inflata 8 0.5 
Thyasira subequatoria 14 180 
Thyasira subovata subovata 18 Ne 
Thyasira transversa 4 0.3 
Thyasira sp. 17 1 0.07 
Kelliella atlantica 30 2.0 
Verticordia sp. 240 1 0.07 
Policordia gemma 3 0.2 
Protocuspidaria sp. 256 1 0.07 
Cuspidaria sp. 1 0.07 
Myonera demistriata 8 0.5 
Myonera sp. 1 0.07 
Incerte cedis sp. 256 1 0.07 

Sta. 242, 4,402 m, 38°16.9'S, 51°56.1’W 
Pristigloma alba 2 0:3 
Neilonella whoii 1 0.1 
Pseudotindaria championi 3 0.4 
Prelametila clarkei 29 3.0 


(continues) 


120 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Ledella aberrenta 2 0.3 
Ledella pustulosa argentinea 104 13.6 
Ledella ultima 3 0.4 
Spinula hilleri E 0.7 
Spinula scheltemae 68 8.8 
Propeleda louiseae 25 9:3 
Yoldiella americana Я 0.9 
Yoldiella blanda 119 15.4 
Yoldiella fabula 8 Mi] 
Yoldiella inconspicua profundorum 50 6.5 
Silicula mcalesteri 1 0.1 
Malletia abyssorum 23 350 

- Malletia grasslei 15 2.0 
Thyasira ferruginea 28 3.7 
Thyasira inflata 4 0.5 
Thyasira subequatoria 11 1.4 
Thyasira transversa 2 08 
Thyasira sp. 17 3 0.4 
Thyasira sp. 21 242 SO 
Kelliella atlantica 1 0.1 
Cuspidaria circinata 3 0.4 
Cuspidaria sp. 242 1 0.1 
Tropidomya sp. 3 0.4 

Sta. 252, 4,435 т, 38°29.8'S, 52°09.1 W 
Pseudotindaria championi 2 12 
Ledella aberrenta 4 2.4 
Yoldiella americana 1 0.6 
Yoldiella blanda 42 25.6 
Yoldiella inconspicua profundorum 23 14.0 
Malletia abyssorum 86 52.4 
Thyasira ferruginea 6 3.6 

Sta. 247, 5,223 m, 43°33.0’S, 48°58.1’W 
Neilonella whoii 6 acd) 
Prelametila clarkei 239 19.1 
Phaseolus sp. c 426 34 
Ledella aberrenta 34 DET. 
Ledella pustulosa argentinea DT 4.5 
Spinula hilleri 24 1.9 
Propeleda louiseae 2 0.2 
Yoldiella americana 293 23.4 
Yoldiella blanda 106 Oro 
Yoldiella fabula 16 19 
Yoldiella inconspicua profundorum 1 0.08 
Thyasira transversa 6 0.5 
Thyasira зр. 8 a 12 1.0 
Cuspidaria sp. 31 219 

WEST EUROPEAN BASIN 

Sta. S29, 119 m, 47°40.0’N, 05°00.0’W 
Nuculana commutata 16 22.9 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Pectinidae sp. c 
Similipecten similis 
Anomiidae sp. 
Thyasira croulinensis 
Kelliella miliaris 
Carditidae sp. 2 
Astarte sp. 5 
Cardiidae sp. 6 
Corbula sp. 2 
Pandora pinna 
Policordia insoleta 
Cuspidaria sp. 
Sta. Biogas | DS01, 400 т, 47°56.5'N, 
07 °40.2W 
Pseudamussium clavatum 
Tridonta elliptica 


Sta. 309, 485 т, 52°21.1’N, 12°07.4W 
Yoldiella frigida 
Limatula bisecta 
Flexopecten proteus 
Similipecten similis 
Delectopecten vitreus 
Thyasira croulinensis 
Thyasira ferruginea 
Thyasira pygmaea 
Thyasira sp. 309 
Mysella verrilli 
Carditidae sp. 2 
Astarte sp. 1 
Astarte sp. 3 
Kelliella biscayensis 
Verticordia sp. 309 
Cuspidaria cuspidata 
Cuspidaria sp. 3 
Cardiomya costellata 
Sta. Incal 0$03, 609 m, 57°25.5’М, 
11°034’W 
Ledella pustulosa pustulosa 
Yoldiella frigida 
Yoldiella jeffreysi 
Limopsis aurita 


Sta. пса! 0$04, 619 m, 57°23.2’N, 
11°06.5'W 
Ledella pustulosa pustulosa 
Yoldiella frigida 
Yoldiella lucida 
Bathyarca pectunculoides 
Limopsis aurita 
Limea sarsi 


No. of 
Individuals 


— — 
= = © = OO N BR © © © © — 


124 


Occurrence 
in % 


(continues) 


122 ALLEN 


(continued) 


No. of Occurrence 
Sample Individuals in % 
Musculus marmoratus 3 $22 
Dacrydium sp. 1 0,4 
Thyasira succisa succisa 24 9.7 
Tridonta elliptica 6 2.4 
Acanthocardia echinata 2 0.8 
Ciliatocardium ciliatum 3+1s 12 
Kelliella atlantica 1 0.4 
Cochlodesma tenerum 4 1.6 
Thracia sp. 2 0.8 
Lyonsiella abyssicola 1 0.4 
Cuspidaria parva 4 126 
Cuspidaria sp. 2 27 12.9 
Sta. S56, 641 m, 43°43.0’N, 03°47.8’W 
Solemya sp. 1 0.05 
Neilonella salicensis 3 0.2 
Ledella acuminata 1 0.05 
Limatula subovata 1 0.05 
Delectopecten sp. a 19 1.0 
Thyasira croulinensis 151 1.9 
Thyasira equalis 15 0.8 
Thyasira eumyaria 106 5.6 
Thyasira ferruginea 2 0.1 
Thyasira succisa succisa 86 4.5 
Thyasira sp. (robusta) 4 0.2 
Montacuta ovata 1 0.05 
Kelliella biscayensis 150% 19: 
Thracia pubescens 3 OZ 
Xylophaga sp. 1 0.05 
Cuspidaria parva 3 0.2 
Cuspidaria sp. 56 1 0.05 
Rhinoclama halimera 1 0.05 
Lyonsiella formosa 3 02 
Incerte cedis sp. 72 1 0.05 
Sta. S40, 860 m, 43°35.6’N, 03°24.8’W 
Nuculoidea bushae 1 0.07 
Thyasira croulinensis 8 0.6 
Thyasira equalis 7 0.5 
Thyasira ferruginea 1 0.07 
Thyasira obsoleta 69 4.9 
Thyasira pygmaea > 0.2 
Thyasira transversa 16 1.1 
Thyasira sp. 40 2 0.1 
Kelliella biscayensis 1300 91.8 
Thracia gracilis 1 0.07 
Thracia pubescens 8 0.6 
Sta. 314, 1,015 m, 51°54.7’N, 12°27.3’W 
Microgloma turnerae 3 0.1 
Nuculoidea bushae 42 1.4 
Ledella pustulosa pustulosa 1 0.03 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 123 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Yoldiella jeffreysi 2 0.07 
Yoldiella sp. 2 0.07 
Silicula filatovae 1 0.03 
Limopsis cristata agg. 1 0.03 
Delectopecten vitreus 3 0.1 
Thyasira eumyaria 4 0.1 
Thyasira sp. 314 1 0.03 
Mysella verrilli 4 0.1 
Abra profundorum 15 0.5 
Kelliella biscayensis 2900 95.7 
Carditidae sp. 2 2 0.07 
Cuspidaria parva 2 0.07 
Incerte cedis sp. 314 2 0.07 


Sta. S63, 1,336 m, 46°17.5'N, 04°45.2°W 


Nuculoma granulosa a 23.3 
Ledella acuminata 1 0.8 
Ledella pustulosa pustulosa 1 0.8 
Ledella similis 8 6.0 
Nuculana vestita 1 0.8 
Dacrydium wareni 4 3.0 
Parvamussium sp. a 2 15 
Delectopecten sp. a 40 30.1 
Thyasira equalis 1 0.8 
Thyasira obsoleta 8 6.0 
Thyasira succisa atlantica 12 9.0 
Thyasira succisa succisa 14 10:5 
Cardiomya costellata 2 1.5 
Rhinoclama notabilis 1 0.8 
Halonympha atlanta 1 0.8 
Halonympha depressa 4 3.0 
Lyonsiella abyssicola 2 ie 
Sta. S66, 1,472 т, 46°16.3’М, 04°44.0’W 
Bathypecten sp. c 1 33.9 
Cyclopecten sp. a 1 33-3 
Thyasira platyssima 1 eh 
Sta: 313, 1/500 m, 51, 32.2'N, 12°35. 9 W 
Microgloma turnerae © 0.07 
Deminucula atacellana 15 0.4 
Nuculoidea bushae 46 La 
Neilonella salicensis 432 10.4 
Ledella acuminata 1 0.08 
Ledella pustulosa pustulosa 456 10.9 
Yoldiella curta 106 2:5 
Yoldiella pseudolata 457 TTC 
Limopsis cristata cristata 1850 44.4 
Dacrydium ockelmanni 10 072 
Limatula louiseae 2 0.05 
Limatula subovata 356 8.5 
Bathypecten eucymatus 16 0.4 


(continues) 


124 


ALLEN 


(continued) 


Sample 


Thyasira croulinensis 
Thyasira equalis 
Thyasira eumyaria 
Thyasira ferruginea 
Thyasira obsoleta 
Thyasira sp. 1 
Thyasira sp. 2 a 
Thyasira sp. 15 
Thyasira sp. 17 
Mysella verrilli 
Mysella sp. 1 
Kelliella atlantica 
Lyonsiella perplexa 
Incerte cedis sp. 313 


Sta. 544, 1,739 m, 43°40.8’М, 03°35.2W 
Microgloma turnerae 
Deminucula atacellana 
Nuculoidea bushae 
Tindaria hessleri 
Neilonella salicensis 
Spinula filatovae 
Yoldiella curta 
Yoldiella inconspicua inconspicua 
Yoldiella pseudolata 
Portlandia lenticula 
Malletia johnsoni 
Limopsis aurita 
Dacrydium ockelmanni 
Bathypecten sp. c 
Cyclopecten sp. a 
Thyasira brevis 
Thyasira croulinensis 
Thyasira equalis 
Thyasira eumyaria 
Thyasira ferruginea 
Thyasira obsoleta 
Thyasira tortuosa 
Thyasira sp. 15 
Thyasira sp. 45 
Kelliella atlantica 
Myonera limatula 


Sta. Biogas Ш DS49, 1,865 m, 44°05.9’N, 
04°15.9’W 

Deminucula atacellana 

Nuculoidea bushae 

Neilonella salicensis 

Ledella pustulosa pustulosa 

Yoldiella curta 

Yoldiella lata 

Malletia johnsoni 


No. of 
Individuals 


4 
313 


— 


La 
WA © © — O> = WN O1 © 00 


— 


Occurrence 
in % 


DONOSO NDS A i ae te ee 
OA BRANWBANOANA-ABD = © O1 O1 O1 1 © O1 O OU 


2 


9. 
6. 
de 
4. 
le 
99: 
0. 


© = © = N N © 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 125 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Thyasira obsoleta 1 0.3 
Thracia pubescens 1 0.3 
Policordia gemma 1 08 
Cuspidaria parva 1 0.3 
Incerte cedis 1 0:3 

Sta. Biogas VI DS88, 1,894 m, 44°05.2’N, 

04°15.7’W 
Deminucula atacellana 7 3.4 
Nuculoidea bushae 14 877. 
Neilonella salicensis > 14.9 
Ledella pustulosa pustulosa % 3.4 
Yoldiella curta 5 2.4 
Yoldiella lata 40 19.2 
Thyasira equalis 3 1.4 
Thyasira obsoleta 3 1.4 
Axinodon symmetros 1 0.5 
Kelliella atlantica 4 1.9 
Cuspidaria parva 1 0.5 
Luzonia simplex 92 44.2 

Sta. Biogas VI DS87, 1,913 m, 44°05.2’N, 

04°19.4’W 
Solemya sp. 1 0.1 
Microgloma turnerae 58 6.4 
Deminucula atacellana 79 8.8 
Nuculoidea bushae 14 1.6 
Neilonella salicensis 178 19.2 
Ledella pustulosa pustulosa 123 TT 
Yoldiella curta 7 0.8 
Yoldiella insculpta 1 0.1 
Yoldiella lata 550 61.1 
Yoldiella obesa incala 5 0.6 
Yoldiella veletta 1 0.1 
Malletia johnsoni 1 0.1 
Limopsis cristata cristata 1 OF 
Dacrydium abyssorum 13 1.4 
Dacrydium ockelmanni 30 oro 
Thyasira brevis 53 5.9 
Thyasira equalis 1 0.1 
Thyasira obsoleta 2 0.2 
Thyasira subcircularis 2 0.2 
Thyasira succisa succisa 1 0.1 
Thyasira ultima 1 0.1 
Thyasira sp. 1 0.1 
Kelliella atlantica 188 20.9 
Cuspidaria parva 42 4.7 

Sta. S65, 1,922 m, 46°15.0'N, 04°35.0’W 
Microgloma turnerae 148 42.8 
Deminucula atacellana 3 0.9 
Nuculoidea bushae 1 0.3 


(continues) 


126 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Neilonella salicensis 2 0.6 
Yoldiella curta 19 95 
Yoldiella insculpta 42 112,1 
Yoldiella pseudolata 25 та 
Silicula fragilis 2 0.6 
Malletia johnsoni 50 14.5 
Hyalopecten parvulinus 8 28 
Parvamussium sp. a 5 1:8 
Cyclopecten sp. a 2 0.6 
Thyasira brevis 15 4.3 
Thyasira sp. 65 15 3.8 
Kelliella atlantica 1 0.3 


Cuspidaria parva 2 
Myonera atlantica 2 
Myonera paucistriata 4 152 
Lyonsiella subquadrata 2 


Sta. Biogas VI DS86, 1,950 m, 44°04.8'N, 


04*18.7"W 
Deminucula atacellana 105 EN 
Neilonella salicensis 199 14.7 
Ledella pustulosa pustulosa 76 5:6 
Yoldiella curta 28 24 
Yoldiella lata 325 23.9 
Yoldiella obesa incala 7 0.5 
Malletia johnsoni 4 0.3 
Dacrydium ockelmanni 36 27 
Limatula subovata 2 0.2 
Thyasira brevis 146 10.8 
Thyasira equalis 4 068 


Thyasira ferruginea 7 
Thyasira obsoleta 9 . 
Thyasira subcircularis 1 0.07 
Thyasira succisa succisa 1 
Thyasira ultima 1 

2 


Mysella tumidula 0.2 
Axinodon symmetros 146 10.8 
Kelliella atlantica 157 11.6 
Policordia gemma 6 0.4 
Cuspidaria parva 58 4.3 
Luzonia simplex 50 uh 

Sta. Biogas IV DS52, 2,006 m, 44°06.3’N, 

04°22.4’W 
Deminucula atacellana 30 4.7 
Nuculoidea bushae 24 3.8 
Tindaria callistiformis 1 02 
Neilonella salicensis 16 25 
Ledella pustulosa pustulosa 47 | 7:3 
Spinula filatovae 1 0.2 
Yoldiella curta 2 0.3 
Yoldiella jeffreysi 1 0.2 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 127 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Yoldiella lata 183 28.6 
Malletia johnsoni 5 0.8 
Dacrydium ockelmanni 19 2.0 
Limatula subovata 1 0.2 
Thyasira brevis 78 V2 
Thyasira equalis 2 0.3 
Thyasira ferruginea 2 3 
Thyasira obsoleta 36 5.6 
Thyasira subcircularis 1 0.2 
Thyasira succisa succisa 1 0:2 
Axinodon symmetros о 0:5 
Kelliella atlantica 158 24.7 
Cuspidaria parva 33 Dz 

Sta. Polygas DS26, 2,076 т, 44°08.2’М, 

0415.0'W 
Deminucula atacellana 12 0.9 
Nuculoidea bushae 11 0.8 
Ledella pustulosa pustulosa 57 4.2 
Yoldiella curta 2 0.2 
Yoldiella lata 1095 81.0 
Yoldiella obesa incala 3 0.2 
Dacrydium abyssorum 9 0.7 
Thyasira brevis 26 1.9 
Thyasira equalis 2 0.2 
Thyasira obsoleta 6 0.4 
Axinodon symmetros 1 0.07 
Kelliella atlantica 123 9.1 
Cuspidaria parva 2 0.4 

Sta. Incal 0302, 2,081 m, 57°58.8'N, 

10°48.5°W 
Microgloma turnerae i 0.2 
Deminucula atacellana 6 0.1 
Nuculoidea bushae 7 0:2 
Neilonella salicensis 91 2.0 
Ledella pustulosa pustulosa 2155 46.0 
Yoldiella curta 544 117 
Yoldiella inconspicua inconspicua 4 0.09 
Yoldiella jeffreysi 67 1.4 
Yoldiella lata 236 Ой 
Yoldiella obesa incala 452 9.8 
Malletia cuneata 62 £33 
Malletia johnsoni 75 10 
Pectinidae sp. 1 O02 
Modiolus sp. 2 0.04 
Musculus discors 1 0.02 
Dacrydium ockelmanni 31 0.7 
Limatula subovata 164 2.0 
Thyasira brevis 5 0.1 
Thyasira ferruginea 16 03 
Thyasira subovata subovata 193 4.2 


(continues) 


128 


ALLEN 


(continued) 


Sample 


Kelliella atlantica 
Mysella verrilli 
Thracia sp. 
Verticordia quadrata 
Verticordia sp. 
Policordia gemma 
Cuspidaria obesa 
Cuspidaria parva 
Cardiomya knudseni 
Incerte cedis 


Sta. Biogas | DS06, 2,090 т, 47°30.5’N, 
08°18.5’W 

Ledella pustulosa pustulosa 

Yoldiella curta 

Yoldiella lata 


Sta. пса! DS01, 2,091 m, 57°59.7’N, 
10°39.8’W 
Microgloma turnerae 
Deminucula atacellana 
Neilonella salicensis 
Ledella pustulosa pustulosa 
Silicula fragilis 
Yoldiella curta 
Yoldiella inconspicua inconspicua 
Yoldiella jeffreysi 
Yoldiella lata 
Yoldiella obesa incala 
Malletia cuneata 
Malletia johnsoni 
Dacrydium ockelmanni 
Limatula subovata 
Thyasira subovata subovata 
Thyasira sp. 1 
Thyasira sp. 2 а 
Kelliella atlantica 
Kelliella miliaris 
Thracia pubescens 
Cochlodesma tenerum 
Verticordia sp. 
Policordia atlantica 
Cuspidaria parva 
Sta. Polygas DS25, 2,096 m, 44°08.2’N, 
04°15.7’W 
Deminucula atacellana 
Nuculoidea bushae 
Neilonella salicensis 
Ledella pustulosa pustulosa 
Yoldiella curta 
Yoldiella lata 


No. of 
Individuals 


so 
1 


Occurrence 


in % 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 129 


(continued) 


No. of Occurrence 
Sample Individuals in % 

Yoldiella obesa incala 3 0.8 
Malletia johnsoni 5 1.4 
Dacrydium ockelmanni 1 0.3 
Thyasira obsoleta 2 0.5 
Thyasira subcircularis 1 0.3 
Kelliella atlantica 51 13.9 
Policordia atlantica 1 0.3 
Cuspidaria parva 5 1.4 
Luzonia simplex 7 1.9 

Sta. Polygas DS17, 2,103 т, 47°32.0’М, 

08°45.5'W 
Yoldiella insculpta 1 8.6 
Dacrydium ockelmanni 2 lee 
Thyasira obsoleta 2 Е 
Cuspidaria parva 13 ae 

Sta. Biogas Ш DS37, 2,110 m, 47°31.8'N, 

08°34.6'W 
Deminucula atacellana 1 2.3 
Yoldiella ella 1 2 
Yoldiella jeffreysi 1 2e 
Yoldiella insculpta 12 Ta 
Yoldiella lata 12 27.3 
Malletia johnsoni > 6.8 
Dacrydium ockelmanni 3 6.8 
Axinulus incrassatus 3 6.8 
Thyasira brevis 1 2,83 
Thyasira obsoleta > 6.8 
Cuspidaria parva 1 2.3 

Sta. Biogas Ш DS50, 2,124 т, 44°08.9’М, 

04°15.9°W 
Deminucula atacellana 1 0:5 
Nuculoidea bushae 3 1.6 
Ledella pustulosa pustulosa 15 7.9 
Yoldiella fabula 1 0.5 
Yoldiella lata 153 80.1 
Dacrydium ockelmanni 7 37 
Thyasira obsoleta 1 0.5 
Kelliella atlantica 8 4.2 
Cuspidaria parva 2 1:1 

Sta. Biogas IV DS63, 2,126 т 47°32.8’М, 

08°35.0' W 
Deminucula atacellana 1 0.6 
Neilonella salicensis 5 270 
Ledella pustulosa pustulosa 78 43.3 
Ledella sublevis 1 0.6 
Yoldiella curta 19 10.6 
Yoldiella insculpta 23 12.8 
Yoldiella jeffreysi 1 0.6 
Dacrydium ockelmanni 6 oo 


(continues) 


130 


ALLEN 


(continued) 


Sample 


Axinulus incrassatus 
Thyasira brevis 
Thyasira obsoleta 
Kelliella atlantica 
Cuspidaria parva 
Tropidomya abbreviata 


Sta. Biogas | DS09, 2,130 m, 47°30.2’N, 
08°16.0’W 

Ledella pustulosa pustulosa 
Sta. Polygas DS18, 2,138 m 47°32'N, 
08°45.5’W 

Deminucula atacellana 

Nuculoidea bushae 

Ledella pustulosa pustulosa 

Yoldiella insculpta 

Yoldiella lata 

Malletia johnsoni 

Dacrydium ockelmanni 

Limatula subovata 

Limatula margaretae 

Thyasira brevis 

Thyasira obsoleta 

Policordia gemma 

Cuspidaria obesa 

Cuspidaria parva 

Incerte cedis 


Sta. Biogas Il DS32, 2,138 m, 47°32.2'N, 
08°05.3’W 
Deminucula atacellana 
Neilonella salicensis 
Ledella aberrata 
Ledella pustulosa pustulosa 
Yoldiella curta 
Yoldiella insculpta 
Yoldiella lata 
Malletia johnsoni 
Dacrydium ockelmanni 
Axinulus incrassatus 
Thyasira equalis 
Thyasira obsoleta 
Thyasira succisa succisa 
Limatula subovata 
Cuspidaria parva 
Sta. Biogas Ш DS38, 2,138 m, 47°32.5’N, 
08°35.8'W 
Tindaria callistiformis 
Neilonella salicensis 
Ledella pustulosa pustulosa 
Ledella sublevis 


No. of 
Individuals 


O1 OD 
=aRAOHNM HH R O1 0 — HOR — 


Occurrence 
in % 


127. 
5.0 
0.6 
0.6 
1,732 
0.6 


100.0 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 131 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Yoldiella curta rs 8.3 
Yoldiella insculpta 2 2.4 
Yoldiella lata 18 155 
Malletia johnsoni 8 9.5 
Dacrydium ockelmanni 1 1.2 
Limatula subovata 2 2.4 
Thyasira brevis 5 6.0 
Thyasira equalis 5 6.0 
Thyasira obsoleta 1 122 
Kelliella atlantica 2 2.4 
Cochlodesma tenerum 1 112 
Cuspidaria parva 1 154 
Cuspidaria sp. 5 6.0 

Sta. Biogas Ш DS36, 2,147 т, 47*32.7'N, 

08°36.5'W 
Ledella pustulosa pustulosa у 18.0 
Yoldiella curta 5 12:8 
Yoldiella insculpta 5 12.8 
Yoldiella lata 7 18.0 
Malletia johnsoni 2 Did 
Dacrydium ockelmanni 1 2.6 
Thyasira brevis 8 20.5 
Thyasira incrassata 4 10.3 

Sta. Biogas V DS70, 2,150 m, 44°08.8’N, 

0417.4'W 
Ledella pustulosa pustulosa i 6.3 
Malletia johnsoni 2 12,5 
Thyasira equalis 2 12:5 
Thyasira ferruginea 4 25:0 
Thyasira subcircularis 1 6.3 
Axinodon symmetros 1 6.3 
Kelliella atlantica 5 Sa 

Sta. Biogas IV DS64, 2,156 m, 47*29.2'N, 

08°30.7’W 
Deminucula atacellana € 2.2 
Ledella pustulosa pustulosa 58 42.3 
Yoldiella curta и 5.1 
Yoldiella insculpta 18 iil 
Yoldiella jeffreysi 2 18 
Yoldiella lata 12 8.8 
Malletia johnsoni 6 4.4 
Dacrydium ockelmanni 14 10.2 
Limatula subovata 5 AR 
Axinulus incrassatus 2 La 
Thyasira brevis 5 Sik 
Thyasira ferruginea 1 0.7 
Thyasira subovata subovata 1 0.7 
Policordia atlantica e 1.5 
Cuspidaria obesa 1 OR; 


(continues) 


132 


ALLEN 


(continued) 


Sample 


Sta. Biogas | DS13, 2,165 m, 47°33.7’N, 
08°35.5’W 

Ledella pustulosa pustulosa 

Spinula subexisa 

Yoldiella curta 

Yoldiella insculpta 

Yoldiella lata 

Portlandia lenticula 

Policordia gemma 

Cuspidaria parva 


Sta. Biogas | DS07, 2,170 m, 47°30.5’N, 
08°15.5 W 

Ledella pustulosa pustulosa 

Yoldiella curta 

Yoldiella lata 


Sta. Biogas IV DS62, 2,175 m, 47°32.8°’N, 


08°40.0’W 
Deminucula atacellana 
Ledella pustulosa pustulosa 
Yoldiella curta 
Yoldiella ella 
Yoldiella insculpta 
Yoldiella jeffreysi 
Yoldiella lata 
Malletia johnsoni 
Dacrydium ockelmanni 
Limatula subovata 
Axinulus incrassatus 
Thyasira brevis 
Thyasira ferruginea 
Thyasira subovata subovata 
Kelliella atlantica 
Policordia atlantica 
Cuspidaria parva 


Sta. Biogas | DS12, 2,180 т, 47°28.5’N, 
08°35:5W 

Ledella pustulosa pustulosa 

Yoldiella biscayensis 

Yoldiella curta 

Yoldiella lata 

Malletia johnsoni 


Sta. Biogas VI DS71, 2,194 m, 47°34.3’N, 


08°33.8'W 
Deminucula atacellana 
Nuculoidea bushae 
Ledella pustulosa pustulosa 
Yoldiella curta 
Yoldiella insculpta 
Yoldiella inconspicua inconspicua 


No. of 
Individuals 


N = Uy 


со — — — — ро 


р aS 


Occurrence 
in % 


97 


fer ee PS Ss 
— 0110010) — > = — 


44.4 
i 
11.1 
224 
da di 


2.8 
0.9 
41.1 
0 
5.6 
0.9 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 135 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Yoldiella jeffreysi 2 1.9 
Yoldiella lata 19 17.8 
Malletia johnsoni 4 37 
Malletia sp. 1 0.9 
Axinulus incrassatus 2 1.9 
Thyasira obsoleta 1 0.9 
Dacrydium ockelmanni р 2.8 
Limatula subovata 6 5:6 
Kelliella atlantica 1 0.9 
Cuspidaria obesa 1 0.9 
Cuspidaria parva 5 4.7 

Sta. Biogas | DS11, 2,205 m, 47°35.5'N, 

08.33./ WW 
Nuculoidea bushae 1 2.0 
Neilonella salicensis 1 2.0 
Ledella pustulosa pustulosa 11 22.0 
Yoldiella insculpta E 18.0 
Yoldiella lata 8 16.0 
Malletia johnsoni 19 38.0 
Dacrydium ockelmanni 1 2.0 

Sta. Biogas | DS08, 2,210 т, 47°27.6'N, 

08°17.0’W 
Yoldiella insculpta 4 66.7 
Yoldiella lata 1 16.7 
Silicula fragilis 1 16.7 

Sta. Biogas Ш DS35, 2,226 m, 47*34.4'N, 

08°40.7’W 
Ledella pustulosa pustulosa 8 13,1 
Yoldiella insculpta 18 29.5 
Yoldiella lata 9 14.8 
Malletia johnsoni 6 9.8 
Limatula subovata 1 1.6 
Axinulus incrassatus 5 8.2 
Thyasira brevis 3 4.9 
Thyasira obsoleta 1 1.6 
Cuspidaria parva 9 14.8 
Cuspidaria sp. 1 1.6 

Sta. Polygas DS15, 2,246 m, 47°35.2'N, 

08°40.1’W 
Ledella pustulosa pustulosa 4 6.5 
Spinula hilleri 1 1:6 
Yoldiella insculpta 5 8.1 
Yoldiella lata 8 12.9 
Malletia johnsoni 8 12.9 
Dacrydium ockelmanni 9 14.5 
Thyasira brevis 1 18 
Policordia gemma 1 1.6 
Cuspidaria jeffreysi 3 4.8 
Cuspidaria parva 15 24.2 


(continues) 


134 


ALLEN 


(continued) 


Sample 


Cuspidaria sp. 
Poromya tornata 


Sta. Biogas IV DS61, 2,250 т, 47°34.7'N, 


08°38.8'W 
Yoldiella curta 
Yoldiella insculpta 
Yoldiella lata 
Malletia cuneata 
Malletia johnsoni 
Dacrydium ockelmanni 
Limatula subovata 
Axinulus incrassatus 
Thyasira brevis 
Thyasira succisa succisa 
Policordia gemma 
Cuspidaria obesa 
Cuspidaria parva 


Sta. Biogas VI DS72, 2,250 т, 47°38.6'N, 


08°36.1' W 
Yoldiella biscayensis 
Yoldiella lata 
Axinulus incrassatus 


Sta. Polygas DS16, 2,325 m, 47°36.1’N, 
08°40.5'W 

Yoldiella insculpta 

Yoldiella lata 

Thyasira incrassata 

Thyasira obsoleta 

Cuspidaria obesa 

Cuspidaria parva 


Sta. Biogas II DS33, 2,338 т, 47°39.7’N, 
08°40.5'W 

Thyasira obsoleta 

Policordia gemma 

Cuspidaria parva 

Cuspidaria sp. 

Halonympha depressa 
Sta. Biogas V DS65, 2,360 m, 47*36.1'N, 
08°40.5’W 

Yoldiella insculpta 

Thyasira incrassata 

Thyasira obsoleta 

Cuspidaria parva 
Sta. 550, 2,397 m, 43°46.7’N, 03°38.0W 

Pristigloma nitens 

Deminucula atacellana 

Neilonella salicensis 

Ledella sublevis 


No. of 
Individuals 


6 
1 


— 
SN =-N RO = BR BR = © © = 


— 


O) = 


Occurrence 
in % 


9:7 
1.6 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 139 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Ledella ultima 12 $ 
Yoldiella inconspicua inconspicua 343 30.07 
Yoldiella jeffreysi 1 0.1 
Yoldiella pseudolata 153 13.7 
Malletia cuneata 4 0.4 
Malletia johnsoni 13 128 
Dacrydium ockelmanni 52 4.7 
Thyasira brevis 3 31 
Thyasira ferruginea 29 2.6 
Thyasira subovata subovata 106 9.5 
Thyasira sp. S50 3 0.3 
Tellinidae sp. f 3 0.3 
Kelliella atlantica 338 30.0 
Bushia sp. 2 0.2 
Luzonia simplex 1 0.1 

Sta. Biogas IV DS51, 2,430 m, 44°11.3’М, 

04°15.4'W 
Pristigloma nitens 1 0.07 
Deminucula atacellana 5 0.4 
Ledella pustulosa pustulosa 3 0.2 
Yoldiella jeffreysi 27 2 
Yoldiella lata 421 34 
Yoldiella obesa incala 1 0.07 
Malletia johnsoni 10 0.7 
Adipicola simpsoni 12 0.9 
Dacrydium wareni 1 0.07 
Thyasira brevis 19 1.4 
Thyasira equalis 3 0.2 
Thyasira ferruginea 1 0.07 
Kelliella atlantica 789 58.3 
Verticordia quadrata 1 0.07 
Policordia gemma 2 0.2 
Cuspidaria parva 21 1.6 
Cuspidaria sp. 1 0.07 
Luzonia simplex 35 2.6 

Sta. 316, 2,493 m, 50°58.7’N, 13°01.6W 
Pristigloma nitens 40 0.6 
Microgloma turnerae 188 21 
Microgloma yongei 1 0.02 
Microgloma sp. $ 1 0.02 
Deminucula atacellana 727 Tie 
Nuculoidea bushae 1 0.2 
Neilonella salicensis 25 0.4 
Ledella pustulosa pustulosa 12 0.2 
Yoldiella curta 91 1.4 
Yoldiella inconspicua inconspicua 20 0.3 
Yoldiella jeffreysi 85 1.3 
Yoldiella pseudolata 1653 258 
Malletia johnsoni 109 he 
Limatula subovata ? 0.1 


(continues) 


136 


ALLEN 


(continued) 


Sample 


Delectopecten vitreus 
Delectopecten sp. a 
Thyasira croulinensis 
Abra profundorum 
Kelliella atlantica 
Thracia sp. 1 
Verticordia sp. 
Policordia densicostata 
Cuspidaria parva 


Sta. пса! DS06, 2,494 т, 56°26.6 N, 


11 105 
Pristigloma nitens 
Microgloma turnerae 
Deminucula atacellana 
Nuculoma granulosa 
Ledella pustulosa marshalli 
Spinula subexisa 
Yoldiella curta 
Yoldiella inconspicua inconspicua 
Yoldiella jeffreysi 
Yoldiella lata 
Yoldiella sp. 
Malletia cuneata 
Malletia johnsoni 
Bathyarca inaequisculpta 
Limatula subovata 
Thyasira ferruginea 
Thyasira subovata subovata 
Thyasira sp. 
Kelliella atlantica 
Abra profundorum 
Verticordia quadrata 
Cuspidaria parva 
Incerte cedis 


Sta. пса! DSO5, 2,503 т, 56°28.1’N, 


LAIT 
Pristigloma nitens 
Deminucula atacellana 
Neilonella salicensis 
Ledella pustulosa marshalli 
Ledella pustulosa pustulosa 
Spinula subexisa 
Yoldiella biscayensis 
Yoldiella curta 
Yoldiella fabula 
Yoldiella inconspicua inconspicua 
Yoldiella jeffreysi 
Yoldiella lata 
Malletia cuneata 
Malletia johnsoni 


No. of 
Individuals 


CO 
= N 2 OAWNWwWAA 


Occurrence 
in % 


0.03 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 137 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Limatula subovata 7 0.7 
Thyasira subovata subovata 67 6.9 
Kelliella atlantica 310 >18 
Policordia densicostata 8 0.8 

Sta. 916, 25051, 9030.27 3 N, 

13°20.9°W 
Pristigloma nitens 85 12 
Microgloma turnerae 12 0.2 
Deminucula atacellana 22 0.3 
Brevinucula verrilli 2 0.03 
Ledella pustulosa marshalli 6 0.08 
Ledella pustulosa pustulosa 25 0.4 
Spinula subexisa 5 007 
Yoldiella inconspicua inconspicua 300 4.7 
Yoldiella jeffreysi 826 US 
Yoldiella lata 258 3.6 
Yoldiella obesa incala 3 0.04 
Malletia johnsoni 24 003 
Dacrydium ockelmanni 1 0.01 
Limatula margaretae 3 0.04 
Limatula subovata 13 0.2 
Axinulus incrassatus 5 0.07 
Thyasira brevis 1 0.01 
Thyasira croulinensis 7 0.1 
Thyasira ferruginea 8 0.1 
Thyasira obsoleta 1 0.01 
Thyasira subovata subovata 104 475 
Thyasira sp. 1 3 0.04 
Thyasira sp. 2 a a 0.03 
Thyasira sp. 17 4 0.06 
Thyasira sp. 32 1 0.01 
Thyasira sp. 318 3 0.04 
Mysella verrilli 3 0.04 
Mysella sp. 1 2 0.03 
Kelliella atlantica 5445 Tout 
Cuspidaria parva 11 Dé 
Cardiomya costellata 14 0.2 
Rhinoclama abrupta 1 0.01 
Halonympha atlanta 1 0.01 
Protocuspidaria simplis 4 0.06 
Incerte cedis sp. 1 a 1 0.01 
Incerte cedis sp. 318 2 0.03 

Sta. Ch. 10, 2,540 m, 56°37.0’N, 

11%04.0'W 
Pristigloma alba 9 0.1 
Pristigloma nitens 285 49 
Microgloma turnerae 11 0.3 
Deminucula atacellana 147 3.9 
Ledella acuminata 245 6.5 
Spinula subexisa 13 0.3 


(continues) 


138 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Yoldiella dissimilis 9 0.2 
Yoldiella jeffreysi 355 9.4 
Yoldiellas subequilateria 194 501 
Silicula fragilis 7 0.2 
Malletia cuneata 74 1.9 
Malletia obtusa 270 EN 
Limatula subovata 9 0.2 
Cyclopecten ambiannulatus 6 0.2 
Thyasira brevis 3 0.08 
Thyasira croulinensis 12 0.3 
Thyasira subovata subovata 388 108 
Epilepton subtrigonum 5 0.1 
Axinodon symmetros oy 10 
Kelliella atlantica 17722 45.6 
Verticordia triangularis 1 0.03 
Cuspidaria parva 8 02 
Cuspidaria sp. 1 0.03 
Incerte cedis sp. 10 1 0.03 

Sta. Incal DS10, 2,719 m, 50°12.7’N, 

13°16.6'W 
Pristigloma nitens 3 172 
Spinula subexisa 4 dG: 
Yoldiella inconspicua inconspicua 21 8.2 
Yoldiella jeffreysi 48 17.9 
Malletia cuneata 2 0.8 
Malletia johnsoni 3 1.3 
Limopsis cristata agg. 1 0.4 
Dacrydium sp. 1 0.4 
Limatula subovata 1 0.4 
Thyasira brevis 8 3A 
Thyasira ferruginea 1 0.4 
Thyasira subovata subovata 6 2.3 
Kelliella atlantica 156 60.7 
Poromya granulata 2 0.8 

Sta. Biogas IV DS58, 2,775 m, 47°34.1’N, 

09°08.2’W 
Ledella pustulosa marshalli 5 9.8 
Yoldiella jeffreysi 10 19.6 
Malletia cuneata ©) 9.9 
Malletia johnsoni 1 2.0 
Dacrydium sandersi 6 11.8 
Axinulus incrassatus 11 21.6 
Thyasira brevis 4 7.8 
Thyasira ferruginea 2 39 
Thyasira sp. 1 2.0 
Policordia atlantica 1 2.0 
Cuspidaria parva 2 3.9 
Myonera angularis 5 9.8 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 139 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Sta. Biogas VI DS74, 2,777 m, 47°33.0’N, 

09°07.8’W 
Ledella pustulosa marshalli 1+1s 1:5 
Ledella pustulosa pustulosa 1s - 
Yoldiella biscayensis 7 10.5 
Yoldiella jeffreysi 2 3.0 
Yoldiella obesa incala й 10.5 
Dacrydium sandersi 1% 25.4 
Axinulus incrassatus 3 4.5 
Thyasira brevis 1 1.5 
Thyasira subovata subovata 1 1.5 
Policordia gemma 1 1. 
Cuspidaria parva Zr 40.3 

Sta. Biogas IV DS59, 2,790 m, 47°31.7’N, 

09°06.2W 
Ledella pustulosa pustulosa 4 4,7 
Yoldiella biscayensis 4 4.7 
Yoldiella lata 2+2s 2 
Yoldiella obesa incala 3 35 
Malletia abyssorum 1 12 
Malletia polita 1 152 
Dacrydium sandersi 2 2,3 
Axinula incrassata 1 1:2 
Thyasira subovata subovata 2 215 
Thyasira succisa succisa 1 1x2 
Kelliella atlantica 2 2.3 
Cuspidaria obesa 56 65.1 
Cuspidaria sp. 1 $52 

Sta. Biogas VI DS73, 2,805 m, 47*32.1'N, 

09°0.0’W 
Ledella pustulosa marshalli 1 Sore 
Ledella sublevis 1 Soo 
Malletia sp. 1 O 

Sta. Biogas II DS31, 2,813 m, 47*32.5'N, 

09°04.2’W 
Deminucula atacellana 1 0.4 
Yoldiella biscayensis Mi 4.3 
Yoldiella jeffreysi 11 4.3 
Yoldiella insculpta 4 1.6 
Yoldiella obesa incala 2 0.8 
Dacrydium ockelmanni 4 1.6 
Dacrydium sandersi 10 3.9 
Axinula incrassata = 2 
Thyasira equalis 2 0.8 
Thyasira obsoleta 1 0.4 
Thyasira subovata subovata 1 0.4 
Thyasira succisa succisa 1 0.4 
Kelliella atlantica 1 0.4 


(continues) 


140 


ALLEN 


(continued) 


Sample 


Cuspidaria obesa 
Myonera angularis 


Sta. 321, 2,868 m, 50°12.3’N, 13°35.8’W 


Pristigloma nitens 
Microgloma turnerae 
Deminucula atacellana 
Brevinucula verrilli 
Pseudotindaria sp. 
Lametila abyssorum 
Ledella pustulosa marshalli 
Ledella sublevis 
Spinula subexisa 
Yoldiella biscayensis 
Yoldiella jeffreysi 
Malletia cuneata 
Dacrydium ockelmanni 
Limatula subovata 
Parvamussium sp. b 
Bathypecten eucymatus 
Thyasira ferruginea 
Thyasira obsoleta 
Thyasira succisa succisa 
Kelliella atlantica 
Cuspidaria atlantica 
Cuspidaria obesa 


Sta. пса! DSO7, 2,884 m, 55°00.7’N, 
12°31.0'W 


Microgloma turnerae 
Deminucula atacellana 
Nuculanidae sp. 

Ledella pustulosa marshalli 
Spinula subexisa 

Yoldiella biscayensis 
Yoldiella jeffreysi 

Malletia cuneata 

Malletia johnsoni 
Bathyarca inaequisculpta 
Limatula subovata 
Dacrydium ockelmanni 
Thyasira subovata subovata 
Thyasira sp. 

Kelliella atlantica 
Cuspidaria parva 
Cuspidaria sp. 

Myonera atlantica 


За. пса! DS08, 2,891 m, 55°02.0'N, 
12°34.6’W 


Deminucula atacellana 
Ledella pustulosa marshalli 
Spinula subexisa 


No. of 
Individuals 


199 


Occurrence 
in % 


17.4 
2.3 


9270 


0.06 


0.06 
28.2 
0.06 
0.06 


30.6 


9.8 
0.06 
10.8 

05 

0.2 
0.06 


083 
11820 
0.8 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 141 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Yoldiella jeffreysi 32 8.6 
Yoldiella inconspicua inconspicua 58 15.6 
Silicula fragilis 2 0.5 
Malletia cuneata 56 15.1 
Malletia johnsoni 2 0.5 
Limatula louiseae 1 0.3 
Limatula subovata 2 0.5 
Thyasira subovata subovata 128 34.4 
Kelliella atlantica 28 7 
Incerte cedis sp. 1 1 0.3 
Incerte cedis sp. 2 1 0.3 

Sta. пса! 0309, 2,897 т, 55°07.7°N, 

12°5 2.6'W 


Pristigloma nitens 1 
Pristigloma alba 3 
Microgloma turnerae a 
Deminucula atacellana 64 27 
Nuculoma granulosa | 3 
1 
1 


Ledella pustulosa pustulosa 

Spinula subexisa 1 0.5 
Yoldiella fabula 3 0.1 
Yoldiella jeffreysi 274 10.4 
Yoldiella inconspicua inconspicua 267 36.6 
Malletia cuneata 1043 44.0 
Malletia johnsoni 16 E 
Bathyarca inaequisculpta 1 0.04 
Limopsis subovata 2 0.08 
Thyasira brevis 19 0.8 
Thyasira equalis 2 0.08 
Epilepton sp. 1 0.04 
Montacutidae sp. T 0.3 
Kelliella atlantica 219 9.2 
Verticordia triangularis 1 0.04 
Cuspidaria parva 1 0.04 
Cardiomya costellata 14 0.6 
Myonera demistriata 10 0.4 

Sta. Ch. 6, 2,900 т, 55°03.0’М, 12°29.0°W 

Pristigloma nitens 416 3.0 
Deminucula atacellana 456 3.6 
Ledella acuminata 6583 47.0 
Spinula subexisa US 0.8 
Yoldiella dissimilis 10 0.07 
Yoldiella lucida 1 0.01 
Yoldiella jeffreysi 6 0.04 
Yoldiella subequilateria 1390 9.9 
Silicula fragilis 23 0.2 
Malletia cuneata 4301 ACT 
Malletia obtusa 40 0.3 
Limopsis tenella À 0.03 
Limatula subovata 9 0.06 


(continues) 


142 


ALLEN 


(continued) 


Sample 


Thyasira brevis 

Thyasira croulinensis 
Thyasira subovata subovata 
Axinodon symmetros 
Kelliella atlantica 

Verticordia triangularis 
Policordia jeffreysi 


Sta. Biogas IV DS57, 2,906 m, 47°30.8'N, 
09°07.6'W 


Ledella pustulosa marshalli 
Ledella sublevis 

Yoldiella fabula 

Malletia cuneata 

Limatula margaretae 
Cuspidaria parva 

Myonera angularis 
Rhinoclama notabilis 


Sta. Biogas VI DS75, 3,250 m, 47°28.1'N, 
09°07.6'W 


Ledella ultima 

Yoldiella biscayensis 

Yoldiella ella 

Yoldiella inconspicua inconspicua 
Yoldiella subcircularis 

Malletia cuneata 

Dacrydium abyssorum 

Kelliella atlantica 

Abra profundorum 


5400925, 3.936 m, oO 06.3'N, 1355.7 


Pristigloma nitens 
Brevinucula verrilli 
Neilonella whoii 

Lametila abyssorum 
Ledella pustulosa marshalli 
Ledella sublevis 

Ledella ultima 

Spinula hilleri 

Yoldiella biscayensis 
Yoldiella ella 

Yoldiella fabula 

Yoldiella inconspicua inconspicua 
Yoldiella jeffreysi 
Malletia cuneata 
Bathyarca inaequisculpta 
Limatula celtica 
Parvamussium permirum 
Cyclopecten sp. a 
Thyasira brevis 

Thyasira transversa 
Thyasira sp. 1 


No. of 
Individuals 


3 
oH 


N & © = = NN = 


SSG ess st eS GOES 


O0 
SJ 


N O) u ok 
DODOO=00h.400 W 


ex ashy 
0 ON = O1NMN 0 


Occurrence 
in % 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Thyasira sp. 15 
Thyasira sp. 17 
Thyasira sp. 30 
Thyasira sp. 323 
Abra profundorum 
Kelliella sp. 323 
Verticordia sp. 
Cetoconcha sp. 
Cuspidaria sp. 
Poromya sp. 
Incerte cedis sp. 323 


Sta. Biogas Ш DS40, 3,345 т, 47°36.4'N, 


09°04.2’W 
Kelliella atlantica 


Sta. Biogas V DS66, 3,480 m, 47°28.2’М, 


09°00.0’W 
Tindaria callistiformis 
Neilonella whoii 
Ledella pustulosa marshalli 
Ledella sublevis 
Ledella ultima 
Yoldiella biscayensis 
Yoldiella ella 
Yoldiella fabula 
Yoldiella inconspicua inconspicua 
Silicula filatovae 
Malletia cuneata 
Limatula margaretae 
Thyasira ferruginea 
Kelliella atlantica 
Policordia gemma 


Sta. Biogas Ш DS41, 3,548 m, 47°28.3'N, 


09%07.2W 
Deminucula atacellana 
Neilonella whoii 
Ledella pustulosa hampsoni 
Ledella pustulosa pustulosa 
Ledella sublevis 
Spinula subexisa 
Yoldiella biscayensis 
Yoldiella fabula 
Yoldiella inconspicua inconspicua 
Yoldiella jeffreysi 
Malletia cuneata 
Dacrydium sandersi 
Limatula margaretae 
Axinulus incrassatus 
Thyasira subovata subovata 
Cuspidaria obesa 


No. of 
Individuals 


pores AS ESE ES ONES (AS 


NO © 
NN SS SS Oo) ISO) Goes EN IN N Sy EN SY NS) 


— 


— 


Occurrence 
in % 


0.4 
1.0 
0.4 


100.0 


N © 


— 


N 
SSN SS (59) [ROY © se N N EN NN EBS | 
ODORS ONDWAOONWOO0O0 OO 


(continues) 


143 


144 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Sta. Biogas IV DS60, 3,742 m, 47°26.8'N, 

09°07.2’W 
Neilonella whoii 2 3.6 
Ledella sublevis 6 10.9 
Yoldiella biscayensis 24 43.6 
Yoldiella ella 6 10.9 
Malletia cuneata 10 18.2 
Limatula margaretae 1 18 
Thyasira ferruginea 1 1.8 
Abra profundorum 2 3.8 
Verticordia triangularis 1 1.8 
Cuspidaria parva 2 3.6 

Sta. 325, 3,846 т, 50°06.2’N, 14°20.9W 
Protocuspidaria verityi 1 20.0 
Cuspidaria sp. 1 1 20.0 
Rhinoclama notabilis 3 60.0 

Sta. 326, 3,859 m, 50°04.9’М, 14°23.8 W 
Pristigloma nitens 4 1.7 
Neilonella whoii 5 22 
Lametila abyssorum 2 0.9 
Ledella pustulosa pustulosa 12 542 
Ledella sublevis 12 52 
Ledella ultima 7 3.0 
Silicula filatovae 1 0.4 
Yoldiella biscayensis 29 12:3 
Yoldiella ella 12 5.2 
Yoldiella inconspicua inconspicua 3 188 
Malletia cuneata 85 36.6 
Limatula margaretae 3 43 
Thyasira croulinensis 1 0.4 
Thyasira transversa 12 SZ 
Thyasira sp. 17 6 2.6 
Kelliella atlantica 33 14.2 
Cuspidaria parva 1 0.4 
Cuspidaria sp. 326 4 dd 

Sta. Biogas III DS44, 3,992 m, 47°33.2’N, 

09°42.0’W 


Yoldiella biscayensis 4 
Malletia cuneata 2 
Bathyarca inaequisculpta 41 17.4 
Dacrydium abyssorum 4 
Thyasira equalis 1 
Kelliella atlantica 1 


Sta. Biogas IV DS56, 4,050 m, 47°32.7'N, 
09°28.2’W 

Tindaria callistiformis 

Neilonella whoii 

Ledella ultima 

Yoldiella biscayensis 


= № — 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Yoldiella ella 3 4.1 
Yoldiella fabula 1 1.4 
Malletia cuneata 4 5.4 
Dacrydium abyssorum 15 20.3 
Thyasira ferruginea 1 1.4 
Kelliella atlantica 5 6.8 
Abra profundorum 4 5.4 
Myonera sp. 33 44.6 
Incerte cedis 1 1.4 

Sta. Biogas Ш DS42, 4,104 т, 47*32.1'N, 

09°35.6'W 
Yoldiella ella 1 14.3 
Malletia cuneata 2 28.6 
Dacrydium abyssorum 1 14.3 
Thyasira equalis 1 14.3 
Abra profundorum 2 28.6 

Sta. Biogas И DS30, 4,106 m, 47°38.3N, 

09°33.9’W 
Ledella ultima 1 5.6 
Yoldiella fabula 1 5.6 
Malletia abyssorum 4 22.2 
Malletia cuneata 1 5.6 
Limatula margaretae 3 vo 7 
Thyasira brevis 1 5.6 
Thyasira equalis 3 16.7 
Thyasira ferruginea 1 5.6 
Kelliella atlantica 3 167 

Sta. Biogas IV DS55, 4,125 m, 47°34.9'N, 

09°40.9’W 
Neilonella whoii 3 0.4 
Ledella pustulosa marshalli 2 0.3 
Ledella ultima 12 1.8 
Yoldiella biscayensis 76 ART 
Yoldiella ella 80 11.6 
Yoldiella fabula 3 0.4 
Yoldiella inconspicua inconspicua 58 8.4 
Yoldiella subcircularis de 1.0 
Malletia abyssorum 65 9.5 
Malletia cuneata 109 15.9 
Dacrydium abyssorum 246 35.8 
Limatula margaretae 1 0.2 
Thyasira atlantica 1 02 
Thyasira brevis 3 0.4 
Thyasira equalis 1 0.2 
Thyasira ferruginea 8 1.2 
Kelliella atlantica 137 19.9 
Abra profundorum 10 1.5 
Policordia gemma 9 1:3 
Cuspidaria parva US 22 
Incerte cedis 1 0.2 


(continues) 


145 


146 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Sta. Polygas DS22, 4,144 m, 47°34.1’N, 

09°38.4'W 
Neilonella whoii 2 1.0 
Yoldiella biscayensis 21 10.9 
Yoldiella ella 4 2:1 
Yoldiella inconspicua inconspicua 3 1.6 
Yoldiella subcircularis 4 25 
Malletia abyssorum 12 6.3 
Malletia cuneata 27 14.1 
Dacrydium abyssorum 59 307 
Limatula margaretae 1 0.5 
Thyasira equalis 1 0.5 
Kelliella atlantica 30 18.2 
Kelliella elongata 2 1.0 
Abra profundorum 6 a 
Policordia gemma 4 р 
Cuspidaria sp. 5 2.6 
Rhinoclama notabilis 5 2.6 
Poromya sp. 1 025 

Sta. Biogas V DS67, 4,150 m, 47°31.0’N, 

09°35.0’W 
Pristigloma alba 3 4.4 
Microgloma yongei 2$ - 
Ledella pustulosa marshalli 1 1..5 
Yoldiella biscayensis 4 00 
Yoldiella ella 4 5.8 
Yoldiella inconspicua inconspicua 1 129 
Malletia abyssorum И 10.1 
Malletia cuneata 9 13.0 
Dacrydium abyssorum 23 19.0 
Limatula margaretae 1 45 
Thyasira ferruginea 4 5.8 
Axinodon symmetros 1 1.9 
Kelliella atlantica 6 8.7 
Abra profundorum 2 2.9 
Cuspidaria parva 3 4.4 

Sta. Polygas DS21, 4,190 m, 47°31.5’N, 

09°40.7’W 
Ledella pustulosa marshalli 1 0.8 
Silicula filatovae 1 0.8 
Yoldiella biscayensis 6 4.7 
Yoldiella ella 8 6.3 
Yoldiella fibula 1 0.8 
Yoldiella inconspicua inconspicua 3 2.4 
Yoldiella subcircularis 1 0.8 
Malletia abyssorum 16 12.6 
Malletia cuneata 24 18.9 
Dacrydium abyssorum 26 20.5 
Limatula margaretae 1 0.8 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Thyasira croulinensis 
Kelliella atlantica 
Abra profundorum 
Policordia atlantica 
Cuspidaria parva 
Cuspidaria sp. 
Poromya sp. 

Incerte cedis 


Sta. Biogas Ш DS48, 4,203 m, 44*29.0'N, 


04°54.0’W 
Malletia abyssorum 
Dacrydium abyssorum 
Thyasira brevis 
Thyasira ferruginea 


Sta alical mo io. 4.2 1 47-24 
0939.1'W 
Ledella ultima 
Yoldiella biscayensis 
Yoldiella ella 
Yoldiella subcircularis 
Malletia abyssorum 
Malletia cuneata 
Dacrydium abyssorum 
Limatula celtica 
Limatula louiseae 
Thyasira brevis 
Thyasira ferruginea 
Thyasira subovata subovata 
Kelliella atlantica 
Abra profundorum 
Policordia gemma 
Cuspidaria obesa 
Myonera atlantica 


Sta. Polygas DS20, 4,226 m, 47°33.0'N, 


09°36.7'W 
Neilonella whoii 
Ledella pustulosa marshalli 
Ledella ultima 
Yoldiella biscayensis 
Yoldiella ella 
Yoldiella inconspicua inconspicua 
Yoldiella subcircularis 
Malletia abyssorum 
Malletia cuneata 
Dacrydium abyssorum 
Kelliella atlantica 
Abra profundorum 
Policordia gemma 
Cuspidaria parva 


No. of 
Individuals 


N NN Y 


ASN SN 


OMAN © NN © ND = = 0 00 NN WO © 


Occurrence 
in % 


2.4 
6.3 
3.2 
1.6 
225 
6.3 
0.8 
0.8 


ee LIN Е ЖА 
MOOMNAMNDOO © 


(continues) 


147 


148 ALLEN 


(continued) 


Sample 


Sta. Biogas VI DS76, 4,228 т, 47°34.8'N, 
09°33.3W 

Neilonella whoii 

Serapta sp. 

Ledella pustulosa marshalli 

Ledella ultima 

Yoldiella biscayensis 

Yoldiella ella 

Yoldiella fabula 

Malletia abyssorum 

Malletia cuneata 

Dacrydium abyssorum 

Limatula margaretae 

Thyasira equalis 

Thyasira ferruginea 

Thyasira transversa 

Thyasira sp. 

Kelliella atlantica 

Kelliella sp. 

Abra profundorum 

Policordia sp. 
Sta. Biogas Ш DS47, 4,230 m, 44*26.8'N, 
04°50.7’W 

Neilonella whoii 
Sta. Biogas VI DS77, 4,240 m, 47°31.8'N, 
09°34.6’W 

Neilonella salicensis 

Ledella ultima 

Silicula filatovae 

Yoldiella biscayensis 

Yoldiella ella 

Yoldiella inconspicua inconspicua 

Yoldiella subcircularis 

Malletia abyssorum 

Malletia cuneata 

Dacrydium abyssorum 

Limatula margaretae 

Thyasira ferruginea 

Kelliella atlantica 

Abra profundorum 

Cuspidaria parva 
Sta. Incal DS14, 4,254 m, 47°32.6’N, 
09°35.7’W 

Neilonella whoii 

Ledella ultima 

Silicula filatovae 

Yoldiella biscayensis 

Yoldiella ella 

Yoldiella fabula 

Yoldiella inconspicua inconspicua 


No. of 


Individuals 


№ — 
© LO w 
> © — © = © O1 = © 


dl 
"| 
co 


№ 
al 
© © 20 NDN0O —= © 


NO 


Occurrence 


in % 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 149 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Yoldiella jeffreysi 1 oS 
Yoldiella subcircularis 3 115 
Malletia cuneata 28 14.1 
Bathyarca inaequisculpta 1 0.5 
Dacrydium abyssorum 75 Of aE 
Limatula subovata 2 1.0 
Thyasira subovata subovata Г. 38 
Montacuta sp. 6 3.0 
Kelliella atlantica 1 0.5 
Abra profundorum 4 2.0 
Poromya tornata 1 0:5 
Protocuspidaria sp. 4 2.0 
Cuspidaria parva 8 4.0 
Rhinoclama notabilis 2 1.0 
Myonera atlantica 12 6.0 

Sta. пса! DS16, 4,268 m, 47°29.6'N, 

09°33.4’W 
Deminucula atacellana 1 0.2 
Neilonella whoii 1 0.2 
Ledella ultima 14 3.0 
Spinula subexisa 1 0.2 
Silicula filatovae 1 0.2 
Yoldiella biscayensis 13 2.8 
Yoldiella ella 57 12:3 
Yoldiella fabula 2 0.4 
Yoldiella inconspicua inconspicua 7 1.5 
Yoldiella jeffreysi 1 0.2 
Yoldiella sp. 5 1 
Malletia cuneata 22 4.8 
Malletia polita % 0.7 
Bathyarca inaequisculpta 4 0.9 
Dacrydium abyssorum 123 26.6 
Limatula margaretae 6 2 
Thyasira brevis 2 0.4 
Thyasira sp. 32 6.9 
Kelliella atlantica 77 187 
Abra profundorum 10 bit 
Verticordia quadrata 2 0.4 
Poromya tornata 6 1.8 
Cuspidaria sp. 1 6 ES 
Cuspidaria sp. 2 1 0.2 
Cuspidaria sp. 3 39 8.4 
Rhinoclama notabilis 15 3,8 
Myonera atlantica 1m 2.4 

Sta. Polygas DS28, 4,413 m, 44°23.8'N, 

04°47.5'W 
Neilonella whoii 1 1.9 
Ledella pustulosa marshalli 3 03 
Silicula filatovae 2 3.8 
Yoldiella fabula 1 1.9 


(continues) 


150 ALLEN 


(continued) 


Sample 


Yoldiella inconspicua inconspicua 
Thyasira brevis 

Kelliella atlantica 

Incerte cedis 


Sta. Biogas IV DS53, 4,425 т, 44°30.4'N, 
04°56.3’W 
Neilonella whoii 
Ledella pustulosa marshalli 
Ledella ultima 
Yoldiella biscayensis 
Yoldiella jeffreysi 
Malletia abyssorum 
Bathyarca inaequisculpta 
Thyasira brevis 
Thyasira ferruginea 
Kelliella atlantica 
Kelliella sp. 
Cuspidaria parva 
Incerte cedis 


Sta. 328, 4,435 m, 50°04.7’N, 15°44.8’W 

Pristigloma nitens 

Neilonella whoii 

Ledella ultima 

Yoldiella biscayensis 

Yoldiella ella 

Yoldiella jeffreysi 

Silicula filatovae 

Malletia abyssorum 

Malletia cuneata 

Limatula celtica 

Dacrydium abyssorum 

Limatula margaretae 

Bathypecten sp. b 

Bathypecten sp. c 

Cyclopecten ambiannulatus 

Cyclopecten sp. a 

Thyasira carrozae 

Thyasira sp. 328 

Kelliella atlantica 

Abra profundorum 

Cuspidaria parva 

Incerte cedis sp. 328 
Sta. Biogas VI DS83, 4,453 m, 44°22.4’N, 
04°51.0’W 

Thyasira sp. 
Sta. Biogas VI DS82, 4,462 m, 44°25.4’N, 
04°52.8’W 

Neilonella salicensis 

Neilonella whoii 


No. of 
Individuals 


7 
15 
21 


ie 


Sea 
© Ud 


aA OO © NN = = © © © BB = ANH NON O 


22 


Occurrence 
in % 


1622 
28.3 
39.6 

SE 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Yoldiella biscayensis 

Yoldiella inconspicua inconspicua 

Malletia abyssorum 

Malletia cuneata 

Bathyarca inaequisculpta 

Thyasira brevis 

Thyasira ferruginea 

Kelliella atlantica 

Cuspidaria sp. 
Sta. Biogas VI DS85, 4,462 m, 44°23.2’М, 
04°50.8’W 

Microgloma yongei 

Ledella ultima 

Yoldiella biscayensis 

Malletia abyssorum 

Thyasira brevis 

Thyasira equalis 

Thyasira ferruginea 


Sta. Biogas VI DS84, 4,466 m, 44°30.0’М, 
04°53.9’W 

Ledella ultima 

Yoldiella fabula 

Malletia abyssorum 

Malletia cuneata 

Thyasira ferruginea 


Sta. Biogas V DS69, 4,510 m, 44°21.9N, 
04°52.4’W 
Ledella ultima 
Yoldiella biscayensis 
Silicula filatovae 
Malletia abyssorum 
Dacrydium abyssorum 
Thyasira brevis 
Thyasira ferruginea 
Kelliella atlantica 
Abra profundorum 


Sta. Biogas II] DS46, 4,521 m, 46°28.6’М, 
10%23.0'W 

Ledella ultima 

Malletia abyssorum 

Malletia polita 

Dacrydium abyssorum 

Thyasira brevis 

Abra profundorum 


Sta. Biogas V DS68, 4,550 m, 46°26.7’N, 
10%23.9W 

Ledella aberrata 

Ledella ultima 

Yoldiella biscayensis 


No. of 
Individuals 


27 


DIO GE Seo) 


N NN 


Occurrence 
in % 


(continues) 


ie 


152 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Yoldiella ella 3 9.4 
Yoldiella jeffreysi 1 31 
Yoldiella subcircularis 1 31 
Malletia abyssorum 2 6.3 
Bathyarca inaequisculpta 2 6.3 
Dacrydium abyssorum 10 31:3 
Kelliella atlantica 6 18.8 
Abra profundorum 1 3 
Sta. 330, 4,632 m, 50%43.5'N, 17°51.7’W 
Pristigloma alba 1 0.08 
Microgloma turnerae 1 0.08 
Nucula sp. 330 2 072 
Brevinucula verrilli 29 2.2 
Neilonella whoii 16 NZ 
Pseudotindaria championi > 0.2 
Ledella acinula 3 EZ 
Ledella pustulosa marshalli 1 0.08 
Ledella sublevis 1 0.08 
Ledella ultima 139 10:5 
Yoldiella biscayensis 29 2? 
Yoldiella ella 137 10.4 
Yoldiella fibula 1 0.08 
Yoldiella inconspicua inconspicua 30 293 
Yoldiella jeffreysi 155 11.8 
Yoldiella subcircularis 8 0.6 
Silicula filatovae 1 0.08 
Malletia abyssorum 10 0.8 
Malletia polita 1 0.08 
Bentharca asperula 3 0.2 
Bathyarca inaequisculpta > 0.4 
Limopsis galathea 47 3.6 
Mytilidae sp. 330 3 0.2 
Limatula celtica 4 0.3 
Limatula margaretae 1 0.08 
Pectinidae sp. b 20 1:30 
Parvamussium permirum 31 2.4 
Cyclopecten ambiannulatus 1 0.08 
Thyasira atlantica 5 0.3 
Thyasira biscayensis 9 0.7 
Thyasira brevis 58 4.4 
Thyasira croulinensis 2 0.2 
Thyasira ferruginea 31 2.4 
Thyasira succisa altlantica 12 0.9 
Thyasira transversa 8 0.6 
Thyasira sp. 15 6 0.4 
Thyasira sp. 17 3 0.2 
Thyasira sp. 47 a 1 0.08 
Thyasira sp. 330 4 0.3 
Thyasira sp. 2 0.2 
Mysella sp. 1 8 0.6 
Abra profundorum 27 2.4 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Kelliella abyssicola 
Kelliella atlantica 
Policordia insoleta 
Policordia laevis 
Laevicordia horrida 
Poromya tornata 
Cuspidaria abbreviata 
Cuspidaria atlantica 
Cuspidaria parva 
Cuspidaria sp. d 
Cuspidaria sp. g 
Cuspidaria sp. h 
Cuspidaria sp. 1 
Cuspidaria sp. 3 
Cardiomya costellata 
Rhinoclama notabilis 
Halonympha depressa 
Myonera demistriata 
Edentaria simplis 
Lyonsiella perplexa 
Lyonsiella smidti 
Lyonsiella subquadrata 
Incerte cedis sp. 1 а 
Incerte cedis sp. 330 
Incerte cedis sp. 334 


Sta. Biogas IV DS54, 4,659 m, 46°31.1'N, 


10°29.2’W 

Pristigloma alba 
Neilonella иво! 
Ledella aberrata 
Ledella ultima 
Yoldiella biscayensis 
Yoldiella jeffreysi 
Yoldiella subcircularis 
Malletia abyssorum 
Malletia cuneata 
Dacrydium abyssorum 
Thyasira brevis 
Thyasira inflata 
Thyasira sp. 

Kelliella atlantica 
Abra profundorum 
Cuspidaria parva 
Incerte cedis 


Sta. Biogas VI DS78, 4,706 т, 46°31.2’N, 


10%23.8W 

Tindaria callistiformis 
Tindaria hessleri 
Neilonella whoii 
Ledella aberrata 


No. of 


Individuals 


— 


—-NMONMH-FANOTP O1 = PWH DD A NW ONON — 


A 
|= — © © = = NANOND — 


198 


Occurrence 
in % 


0.08 
13 
0.2 
0.2 
0.8 

0.08 
0.2 
aD 

0.08 
0.4 
0.4 

0.08 
0.2 
0.3 

0.08 
0.4 
03 
1,9 
0.4 
ORS 
0.3 

0.08 
0.2 
0.2 

0.08 


(continues) 


154 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Ledella galatheae 1 0.2 
Ledella ultima 29 6.8 
Yoldiella biscayensis 48 11.2 
Yoldiella ella 3 OA 
Yoldiella jeffreysi 1 0.2 
Yoldiella subcircularis 4 0.9 
Malletia abyssorum 201 47.0 
Malletia cuneata 21 4.9 
Malletia polita 8 1.9 
Dacrydium abyssorum 21 4.9 
Thyasira equalis 2 0.5 
Mysella tumidula 1 0.2 
Kelliella atlantica 83 eve 
Mactridae sp. 2 0.5 
Policordia sp. 1 0.2 

Sta. Biogas VI DS79, 4,715 m, 46°30.4'N, 

10°27 41 WV 
Microgloma turnerae 2 0.4 
Tindaria miniscula 1 02 
Pseudotindaria erebus 3 0.6 
Neilonella whoii 13 2.8 
Ledella aberrata T7 3.6 
Ledella acinula 15 92 
Ledella ultima 21 4.5 
Yoldiella biscayensis 110 299 
Yoldiella ella 9 1.9 
Yoldiella jeffreysi 2 0.4 
Yoldiella obesa incala 1 0.2 
Yoldiella subcircularis 4 0.9 
Malletia abyssorum 129 BESS 
Malletia cuneata 58 1283 
Malletia polita 12 2.5 
Bathyarca inaequisculpta L 15 
Thyasira sp. 1 0.2 
Kelliella atlantica 56 RS 
Abra profundorum 1 0.2 
Cuspidaria sp. 1 1 0.2 
Cuspidaria sp. 2 7 15 
Myonera angularis 2 0.4 

Sta. Biogas VI DS81, 4,715 m, 46°28.3'N, 

10°24.6’W 
Neilonella whoii 1 5.9 
Ledella aberrata 2 MS 
Ledella galathea 1 5.9 
Yoldiella biscayensis 10 58.8 
Yoldiella subcircularis 1 5.9 
Malletia polita 2 11.8 
Dacrydium abyssorum 1 5.9 


(continues) 


(continued) 


BIVALVIA OF THE DEEP ATLANTIC 


No. of 
Sample Individuals 


Sta. Biogas VI DS80, 4,720 т, 46°29.5’N, 

10°29.5'W 
Neilonella salicensis 
Ledella aberrata 
Ledella ultima 
Yoldiella biscayensis 
Yoldiella ella 
Yoldiella obesa incala 
Yoldiella subcircularis 
Malletia abyssorum 
Malletia cuneata 
Malletia polita 
Dacrydium abyssorum 
Thyasira biscayensis 
Kelliella atlantica 
Cuspidaria sp. 

Sta. Polygas DS23 4,734 т, 46°32.8'N, 

10°21.0’W 
Pristigloma nitens 
Neilonella whoii 
Ledella aberrata 
Ledella ultima 
Ledella sp. 
Yoldiella biscayensis 
Yoldiella ella 
Yoldiella fabula 
Yoldiella jeffreysi 
Malletia abyssorum 
Malletia cuneata 
Malletia polita 
Dacrydium abyssorum 
Thyasira brevis 
Thyasira equalis 
Kelliella atlantica 
Myonera atlantica 


Sta. Incal DS11, 4,823 т, 48°18.8’М, 

19. ley 
Neilonella whoii 
Ledella aberrata 
Ledella ultima 
Yoldiella ella 
Yoldiella jeffreysi 
Yoldiella subcircularis 
Dacrydium abyssorum 
Thyasira biscayensis 
Thyasira brevis 
Thyasira obsolete 
Thyasira transversa 


— 


== W 
|]=NDNODNWONOTBNWODW A 


=. 
O) Er EN 


DD—_0o0O9@0—-DDAN-—-0OU010MN 


— N 


NO 


№ 
N O1 


O —= © — © — — WON = 


Occurrence 
in % 


— 


— 
ON ON OENNOCEN 
VDO=0000o0=NnN=0 


Eh 


(continues) 


155 


156 


ALLEN 


(continued) 


Sample 


Abra profundorum 
Protocuspidaria sp. 


CANARIES BASIN 
Sta. D6697, 1,564 т, 27°57.0'N, 


13°46.2’W 
Deminucula atacellana 
Nuculoidea bushae 
Limopsis cristata cristata 
Thyasira obsolete 
Kelliella atlantica 


Sta. D6696, 1,780 т, 28°06.0’N, 


13°28.0’W 
Nuculoidea bushae 
Malletia cuneata 
Bathyarca pectunculoides 
Propeamussium meridionale 
Thyasira obsolete 
Cuspidaria atlantica 
Cuspidaria jeffreysi 
Cuspidaria sp. 
Bidentaria atlantica 
Lyonsiella formosa 
Lyonsiella subquadrata 


Sta. D6701, 1,934 m, 27°45.2’N, 
14°13.0°W 


Deminucula atacellana 
Nuculoidea bushae 
Tindaria agatheda 
Neilonella salicensis 
Yoldiella insculpta 
Yoldiella pseudolata 
Yoldiella veletta 
Dacrydium ockelmanni 
Thyasira biscayensis 
Thyasira obsolete 
Thyasira sp. 67 
Kelliella atlantica 
Cuspidaria parva 


Sta. D6704, 2,129 т, 27°44.9'N, 
14°25.0’W 


Deminucula atacellana 
Nuculoidea bushae 
Brevinucula verrilli 
Tindaria agatheda 
Neilonella salicensis 
Yoldiella insculpta 
Yoldiella jeffreysi 
Portlandia lenticula 
Dacrydium ockelmanni 


No. of 
Individuals 


2 
1 


NO = N © 


М — Юм 


— ASS SS 


— 
O 
— OO = N = © —= © 2 0 YN O) 


Occurrence 


In % 


4.1 
2.0 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Abra profundorum 
Kelliella atlantica 
Protocuspidaria verityi 
Rhinoclama halimera 


Sta. 06709, 2,351 т, 27°29.8'N, 

15220410 
Pristigloma alba 
Microgloma turnerae 
Brevinucula verrilli 
Bentharca nodulosa 
Pectinidae sp. 
Thyasira obsolete 
Abra profundorum 
Cuspidaria parva 
Cuspidaria sp. 
Rhinoclama halimera 
Halonympha depressa 
Protocuspidaria simplis 
Lyonsiella abyssicola 
Lyonsiella formosa 


Sta. D6707, 2,593 m, 27°29.2'М, 
15°265 WV 

Brevinucula verrilli 

Kelliella atlantica 


Sta. 06710, 2,670 m, 27°23.6'N, 
15°39.6 W 
Pristigloma nitens 
Brevinucula verrilli 
Spinula hilleri 
Yoldiella insculpta 
Bentharca nodulosa 
Kelliella atlantica 
Cuspidaria parva 
Lyonsiella abyssicola 
Sta. D6711, 2,988 m, 27°14.9'N, 
1586334 
Pristigloma alba 
Pristigloma nitens 
Brevinucula verrilli 
Neilonella whoii 
Spinula hilleri 
Malletia cuneata 
Bentharca nodulosa 
Limopsis minuta 
Propeamussium centobi 
Thyasira transversa 
Leptonidae sp. w 
Kelliella atlantica 
Cuspidaria teres 


No. of 
Individuals 


182 
1 
1 


— 


EN К А) м TRS 


© N 


— бо 


Ам = CD TER ss 


Occurrence 
in % 


Ons 
61.9 
0.3 
0.3 


(continues) 


157 


158 


ALLEN 


(continued) 


Sample 


Myonera atlantica 
Myonera sp. 
Poromya tornata 


Sta. 06714, 3,301 т, 27°13.0'N, 
15°41.0’W 

Yoldiella insculpta 

Yoldiella subcircularis 

Malletia abyssorum 

Pectinidae sp. a 

Myonera sp. 

Lyonsiella freilei 


SIERRA LEONE BASIN 


Sta. 142, 1,796 m, 10°30.0’N, 17°51.5’W 
Nuculoidea bushae 
Nuculoma perforata 
Neilonella salicensis 
Ledella lusitanensis 
Spinula filatovae 
Yoldiella bilanta 
Yoldiella curta 
Malletia johnsoni 
Dacrydium ockelmanni 
Limatula subovata 
Bathypecten sp. e 
Cyclopecten sp. a 
Thyasira carrozae 
Thyasira equalis 
Thyasira eumyaria 
Thyasira obsoleta 
Thyasira subcircularis 
Thyasira subovata subovata 
Thyasira succisa atlantica 
Thyasira tortuosa 
Thyasira transversa 
Thyasira ultima 
Thyasira sp. 17 
Policordia atlantica 
Cuspidaria parva 
Luzonia simplex 
Myonera atlantica 
Protocuspidaria simplis 
Lyonsiella formosa 


Sta. 138, 1,976 т, 10°36.0’N, 17°52.0’W 
Nuculoma perforata 
Brevinucula verrilli 
Tindaria hessleri 
Yoldiella bilanta 
Malletia johnsoni 
Bathyarca inaequisculpta 


No. of 
Individuals 


1 
1 
1 


SOS "О 


Occurrence 
in % 


и 
NUBRWADRDOWA 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 199 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Thyasira ferruginea 3 6.3 
Kelliella atlantica 4 8:5 
Kelliella elongata 1 Zo 

Sta. 143, 2,095 т 10°35.0’N, 17°44’W 
Nuculoma perforata 18 16.8 
Brevinucula verrilli 4 3.7 
Malletia johnsoni 1 0.9 
Bathyarca inaequisculpta 9 8.4 
Cyclopecten sp. a 1 0.9 
Thyasira ferruginea 65 60.8 
Kelliella atlantica 9 8.4 
Verticordia triangularis 1 0.9 

Sta. 141, 2,131 т, 10°30.0’N, 17°51.5’W 
Nuculoidea bushae 3 an 
Nuculoma perforata 38 46.3 
Brevinucula verrilli 1 1.2 
Tindaria hessleri 3 Phat 
Ledella pustulosa hampsoni 2 2.4 
Yoldiella bilanta 2 2.4 
Malletia johnsoni 7 69 
Thyasira ferruginea 20 24.4 
Kelliella atlantica 8 9.8 


Эа. 1.9, 2,187 т, 1035/0N, 1753.01 


Nuculoma perforata 27 72.4 
Brevinucula verrilli 3 10,3 
Tindaria hessleri 1 3:0 
Yoldiella americana 1 35 
Yoldiella pseudolata 1 oe 
Bathyarca inaequisculpta 2 6.9 
Sta. 145, 2,192 m, 10°36.0’N, 17°49’W 
Microgloma yongei 19 85 
Nuculoma perforata 25 1,2 
Brevinucula verrilli 10 4.5 
Tindaria hessleri 1 0.5 
Ledella pustulosa hampsoni 1 0.5 
Spinula hilleri 1 0,5 
Yoldiella curta 1 (1.8 
Yoldiella inconspicua africana 29 13.0 
Yoldiella veletta 1 0.5 
Portlandia lenticula 1 0.5 
Pectinidae sp. e 1 Os 
Malletia johnsoni 30 135 
Bathyarca inaequisculpta 9 4.0 
Thyasira croulinensis 30 19.8 
Thyasira equalis 2 0.9 
Thyasira ferruginea 2 0.9 
Thyasira subovata subovata 1% 5.8 
Thyasira tortuosa 3 1.4 
Thyasira ultima 38 17.0 


(continues) 


160 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Policordia atlantica 3 1.4 
Cuspidaria parva 1 0.5 
Sta. 144, 2,357 m, 10°36.0’N, 17°49.0’W 
Nuculoma perforata 39 17.0 
Brevinucula verrilli 9 3.9 
Tindaria hessleri T 3 
Ledella pustulosa hampsoni 3 tes 
Yoldiella bilanta 22 9.6 
Malletia johnsoni 3 aS 
Malletia pallida 29 12% 
Dacrydium ockelmanni 2 0.9 
Parvamussium sp. a 1 0.4 
Cyclopecten sp. a 2 0.9 
Thyasira alleni 1 0.4 
Thyasira croulinensis 1 0.4 
Thyasira eumyaria 4 1.8 
Thyasira ferruginea 20 8.7 
Thyasira obsoleta 2 0.9 
Thyasira subcircularis 1 0.4 
Thyasira tortuosa 1 0.4 
Thyasira ultima 9 3.9 
Kelliella atlantica 40 Tio 
Cochlodesma tenerum 1 0.4 
Verticordia quadrata zZ 0.9 
Verticordia sp. 1 0.4 
Protocuspidaria verityi 1 0.4 
Cuspidaria parva 7 31 
Luzonia simplex 9 3.9 
Sta. 146, 2,891 m, 10°39.5’N, 17°44.5’W 
Brevinucula verrilli 4 16.7 
Neilonella whoii 2 8.3 
Nuculana vestita 1 4.2 
Ledella ultima 5 20.8 
Malletia johnsoni 2 8.3 
Malletia pallida 6 2010 
Bathyarca inaequisculpta 1 4.2 
Thyasira ferruginea 1 4.2 
Kelliella abyssicola 1 4.2 
Incerte cedis sp. 146 1 4.2 
Sta. 147, 2,934 т, 10°38.0’N, 17°52.0’W 
Brevinucula verrilli 51 18.8 
Neilonella salicensis 16 5.9 
Neilonella whoii 4 4.8 
Ledella sublevis 27 9.9 
Ledella lusitanensis 18 6.6 
Ledella ultima 29 10.7 
Spinula filatovae 1 0.4 
Nuculana vestita 2 aa 
Yoldiella inconspicua africana 5 1.8 
Malletia johnsoni 38 14.0 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Malletia pallida 
Bathyarca inaequisculpta 
Dacrydium ockelmanni 
Parvamussium sp. a 
Thyasira croulinensis 
Thyasira equalis 
Thyasira ferruginea 
Axinus grandis 
Kelliella abyssicola 
Rhinoclama halimera 
Incerte cedis sp. 1b 


Sta. 148, 3,828 т, 10°37.0’N, 18°14.0’W 
Nuculoidea bushae 
Brevinucula verrilli 
Neilonella whoii 
Pseudotindaria erebus 
Ledella ultima 

Portlandia abyssorum 
Yoldiella ella 

Malletia abyssorum 
Bathyarca inaequisculpta 
Limopsis galathea 
Limopsis tenella 
Dacrydium abyssorum 
Mysella sp. 1 

Cuspidaria barnardi 


Sta. 149, 3,861 т 10°30.0’N, 18°18.0’W 
Nuculoidea bushae 
Brevinucula verrilli 
Neilonella whoii 
Pseudotindaria erebus 
Portlandia abyssorum 
Yoldiella ella 

Yoldiella jeffreysi 
Silicula filatovae 
Malletia abyssorum 
Limopsis tenella 
Dacrydium abyssorum 
Cyclopecten sp. a 
Thyasira croulinensis 
Cuspidaria sp. 
Verticordia quadrata 
GUINEA BASIN 


Sta. Walda DS28, 1,261 m, 04°21.2’N, 
0435.21 

Nuculoidea bushae 

Tindaria callistiformis 

Neilonella whoii 

Ledella acinula 


No. of 
Individuals 


22 


Occurrence 


in % 


(continues) 


161 


162 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Spinula filatovae 49 44.1 
Spinula subexisa Y 6:3 
Malletia pallida 5 4.5 
Thyasira brevis 3 27 
Thyasira eumyaria 1 0.9 
Thyasira ultima 1 0.9 
Rhinoclama sp. 2 1.8 

Sta. Walda DS27, 1,376 m, 03°30.7’N, 

05°31.8’E 
Spinula filatovae 1 ОЕ 
Portlandia lenticula 2 33.3 
Thyasira brevis 1 16.7 
Thyasira ultima 2 33:3 

Sta. Walda DS26, 1,890 m, 03%05.1'N, 

ODE TE 
Yoldiella bilanta 2 100.0 

Sta. Walda DS19, 2,243 m, 03°48.0’S, 

ОЭ Е 
Thyasira equalis 1 100.0 

Sta. Walda DS25, 2,470 m, 02*19.8'N, 

07°49.2’E 
Brevinucula verrilli 60 47.6 
Neilonella salicensis 31 24.6 
Ledella sublevis 34 30.0 
Thyasira brevis 1 0.8 

Sta. Walda DS20, 2,514 т, 02*32.0'S, 

08°18.1’E 
Deminucula atacellana 1 4.4 
Brevinucula verrilli 6 26.1 
Neilonella salicensis A 17.4 
Ledella sublevis 1 4.4 
Ledella ultima 1 4.4 
Yoldiella artipica 3 13.0 
Yoldiella inconspicua africana 5 Zilch 
Abra longicallis 2 8:7 

Sta. Walda DS22, 3,025 m, 00°35.6’S, 

06°49.4’E 
Brevinucula verrilli 5 45.5 
Limopsis tenella 3 РГ: 
Thyasira brevis 1 9.1 
Verticordia subquadrata 1 9.1 
Myonera sp. 1 9.1 

Sta. Walda DS30, 3,109 m, 04°04.1'N, 

03°42.0'E 
Brevinucula verrilli 22 56.4 
Tindaria miniscula 3 TT 
Pseudotindaria erebus 4 10.3 
Yoldiella fabula 3 TT 
Malletia cuneata 7 18.0 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Sta. Walda DS23, 3,138 m, 00°15.3’S, 
05°47 .8'E 

Brevinucula verrilli 

Yoldiella fabula 

Bentharca asperula 

Cuspidaria ventricosa 


Sta. Walda DS29, 3,147 m, 02°57.0'N, 
04°28.1’E 

Deminucula atacellana 

Brevinucula verrilli 

Yoldiella fabula 

Yoldiella subcircularis 


Sta. Walda DS21, 4,019 m, 02°38.2’S, 
05°40.0’E 

Limopsis tenella 
Sta. Walda DS31, 4,279 m, 03°17.5’N, 
02°01.7 © 

Ledella galatheae 

Ledella ultima 

Yoldiella ella 

Limopsis galathea 


ANGOLA BASIN 


Sta. 203, 542 m, 08°46.0’S, 12°47.0’E 
Nuculoidea bushae 
Nuculoma perforata 
Neilonella salicensis 
Nuculana vestita 
Propeleda paucistriata 
Portlandia minuta 
Cyclopecten sp. a 
Thyasira alleni 
Thyasira bushae 
Thyasira carrozae 
Thyasira croulinensis 
Thyasira equalis 
Thyasira obsolete 
Thyasira subovata minuta 
Thyasira succisa atlantica 
Thyasira transversa 
Thyasira sp. 3 
Thyasira sp. 28 
Thyasira sp. 34 
Montacuta ovata 
Mysella мет! 

Mysella sp. 1 
Epilepton sp. 21 
Tellinidae sp. c 
Tellinidae sp. x 
Kelliella atlantica 


No. of 
Individuals 


— № = = 


N = N N 


163 


Occurrence 
In % 


(continues) 


164 


ALLEN 


(continued) 


Sample 


Policordia atlantica 
Tropidomya diagonalis 
Luzonia simplex 
Incerte cedis sp. 203 a 
Incerte cedis sp. 203 b 


Sta. Walda DS07, 1,227 т, 19%57.0'S, 
11 02/0'E 
Tindaria callistiformis 
Limatula smithi 
Limopsis cristata agg. 
Lucinoma filosa 
Thyasira excavata plicata 
Thyasira sp. 
Sta. Walda DS10, 1,432 m, 18°40.0’S, 
10°56.3°E 
Neilonella seguenza 
Neilonella whoii 
Yoldiella capensis 
Yoldiella curta 
Limopsis cristata lanceolata 
Lucinoma filosa 
Thyasira alleni 
Thyasira carrozae 
Thyasira equalis 
Thyasira subcircularis 
Thyasira subovata subovata 
Thyasira tortuosa 
Thyasira transversa 


Sta. Walda DS14, 1,537 т, 11*57.6'S, 
12°54.3’E 
Limopsis cristata lanceolata 
Sta. 202, 1,643 m, 09°05.0'S, 12°17.0°E 
Pristigloma nitens 
Microgloma turnerae 
Deminucula atacellana 
Nuculoma granulose 
Neilonella salicensis 
Ledella pustulosa hampsoni 
Ledella sp. 
Spinula filatovae 
Spinula hilleri 
Yoldiella artipica 
Yoldiella hanna 
Portlandia lenticula 
Dacrydium abyssorum 
Pectinidae sp. d 
Cyclopecten ambiannulatus 
Thyasira brevis 
Thyasira croulinensis 


No. of 
Individuals 


15 


— — — = = à 


SO A МЮ IND IND A 


Occurrence 
in % 


0.3 
0.04 
15.6 
0.02 

0.8 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 165 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Thyasira ferruginea 11 3.0 
Thyasira inflata 1 0.3 
Thyasira subequatoria 5 1.4 
Thyasira succisa atlantica 114 3143 
Thyasira tortuosa 14 3.9 
Thyasira transversa 10 2:8 
Thyasira ultima 46 12.6 
Thyasira sp. 15 8 22 
Thyasira sp. 202 8 2.2 
Epilepton sp. 21 1 0.3 
Mysella sp. 2 22 6.0 


Mysella sp. 3 1 
Kelliella atlantica 34 
Abra profundorum 1 
Thracia sp. 3 1 
Cuspidaria sp. 202 1 
Myonera sp. 2 0.06 
Lyonsiella sp. 2 
Incerte cedis sp. 1 1 
Incerte cedis sp. 2 1 
Incerte cedis sp. 3 1 
Incerte cedis sp. 4 8 


Sta. Walda DS16, 1,787 m, 10°31.0’S, 
1157818 
Yoldiella inconspicua Africana 2 
Yoldiella similes 4 
Thyasira brevis 2 20.0 
Thyasira eumyaria 1 
Thyasira obsolete 1 


Sta. 201, 2,031 т, 09°29.0'S, 11°34.0’Е 


Solemya sp. 259 2 Os 
Microgloma yongei 18 2 
Deminucula atacellana 2 0.3 
Nuculoidea bushae 8 12 
Neilonella salicensis 41 6.2 
Ledella ultima 1 0.2 
Nuculana vestita 1 0.2 
Yoldiella inconspicua Africana 109 16.5 
Silicula fragilis 6 0.9 
Malletia johnsoni 8 1.2 
Bathyarca inaequisculpta 39 5.9 
Thyasira carrozae 1 0.2 
Thyasira croulinensis 24 3,6 
Thyasira ferruginea 3 0.5 
Thyasira obsolete р We 
Thyasira ultima 3 0.5 
Thyasira sp. 17 3 0.5 
Epilepton sp. 21 2 0.3 
Abra profundorum 5 0.8 
Kelliella atlantica 251 37.9 


(continues) 


166 


ALLEN 


(continued) 


Sample 


Protocuspidaria verity! 

Cuspidaria parva 

Luzonia simplex 
Sta. Walda DSO6, 2,745 т, 22°50.2’S, 
= 

Malletia johnsoni 

Thyasira brevis 


Sta. 200, 2,754 т, 09°41.0’S, 10°55.0’E 
Microgloma yongei 
Nuculoidea bushae 
Neilonella whoii 
Pseudotindaria erebus 
Ledella sublaevis 
Ledella ultima 
Yoldiella artipica 
Malletia johnsoni 
Malletia pallida 
Bathyarca inaequisculpta 
Limopsis galathea 
Lucinidae sp. 

Thyasira brevis 
Thyasira croulinensis 
Thyasira equalis 
Thyasira ferruginea 
Thyasira transversa 
Thyasira sp. 15 
Thyasira sp. (indet. crushed) 
Epilepton sp. 21 

Abra profundorum 
Kelliella atlantica 
Verticordia triangularis 
Cuspidaria sp. 519 
Halonympha atlanta 
Policordia sp. 


Sta. Walda DSO5, 2,992 m, 21%45.0'S, 
11078 

Silicula filatovae 

Yoldiella curta 

Thyasira brevis 

Thyasira ferruginea 


За. Walda DS15, 3,367 т, 12°27.2’S, 
1101.54 

Pseudotindaria erebus 

Yoldiella sp. 

Thyasira brevis 

Cuspidaria sp. 


Sta. 199, 3,779 m, 09°47.0’S, 10°29.0’E 
Neilonella whoii 


No. of 
Individuals 


16 
7 


Ex 
— 
- — —= & N о O1 — 


a — => à 


Occurrence 
in % 


Sed. 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 167 


(continued) 
No. of Occurrence 
Sample Individuals in % 

A ee a ee A A a ee 
Ledella ultima 59 fie 
Yoldiella curta 2 2.4 
Yoldiella ella 2 2.4 
Malletia pallida 11 13.4 
Thyasira sp. 15 3 37 
Cuspidaria sp. 2 2.4 

Sta. 195, 3,797 m, 14°49.0’S, 09°56.0’E 
Solemya sp. 195 10 q 
Pristigloma alba 8 0.9 
Pristigloma nitens 9 1.0 
Nuculoidea bushae 20 21 
Neilonella whoii 46 4.9 
Pseudotindaria erebus 148 15:6 
Ledella ultima 355 37.3 
Spinula hilleri 98 10.4 
Yoldiella bilanta 43 4.7 
Yoldiella fibula 7 0.7 
Portlandia abyssorum 48 5.1 
Malletia pallida 58 6.1 
Bathyarca inaequisculpta 40 4.2 
Limopsis tenella 41 4.3 
Cyclopecten sp. a 5 0.5 
Thyasira subequatoria 8 0.9 
Verticordia quadrata 1 0.1 

Sta. Walda DS13, 3,985 m, 14°21.5’S, 

09°46.2’E 
Pseudotindaria erebus 8 59 
Malletia johnsoni 192 85 
Limopsis tenella 14 6.2 
Dacrydium angulare 10 4.4 
Thyasira brevis 1 0.4 
Lyonsiella freilei 1 0.4 

Sta. Walda DS18, 4,079 m, 06°37.4’S, 

08°18.2’E 
Bentharca asperula 1 50.0 
Limopsis tenella 1 50.0 

Sta. Walda DS04, 4,180 m, 21°59.1’S, 

0901:5'E 
Yoldiella capensis 1 50.0 
Yoldiella hanna 1 50.0 

Sta. Walda DS17, 4,223 т, 09°12.0'S, 

10°29.0’E 
Bentharca asperula 4 80.0 
Limopsis tenella 1 20.0 

Sta. Walda DS12, 4,308 m, 17°32.8’S, 

09°28.7’E 
Tindaria miniscula 5 100.0 


(continues) 


168 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 
Sta. 198 4,566 m, 10°29.0’S, 09°04.0’E 
Pristigloma alba 7 0.9 
Pristigloma nitens 5 0.6 
Nuculoidea bushae 16 2.0 
Tindaria miniscula 7 0.9 
Neilonella whoii 20 25 
Ledella ultima 599 74.5 
Spinula hilleri 3 0.4 
Spinula sp. 3 0.4 
Yoldiella ella 4 0.5 
Yoldiella fibula 10 
Yoldiella similes 12 
Silicula filatovae 13 
Malletia pallida 1 
Bentharca asperula 4 
Bathyarca inaequisculpta 20 
Limopsis galathea 36 


Limatula smithi 1 
Parvamussium sp. a 1 
Thyasira brevis 8 
Thyasira inflata 6 
Thyasira transversa 6 
1 
2 
2 
1 


— 


Abra profundorum 
Kelliella atlantica 
Rhinoclama sp. 
Poromya sp. 256 


Sta. 197, 4,596 m, 10°29.0'S, 09°04’E 


DOODODON2D0OAÁNOO 22 
22 © © — © © © = —= O1 O1 O1 O O O1 


Pristigloma alba 1 0.08 
Pristigloma nitens 3 0.3 
Nuculoidea bushae 34 3:0 
Tindaria callistiformis 4 0.3 
Tindaria miniscula 27 2.4 
Neilonella whoii 29 2.2 
Prelametila sp. 247 2 O22 
Ledella ultima 647 56.5 
Spinula hilleri 9 0.8 
Yoldiella ella 2 072 
Yoldiella fibula 5 0.4 
Yoldiella jeffreysi 39 3.1 
Yoldiella similes 30 =: 
Silicula filatovae 25 22 
Malletia pallida 3 0,8 
Bentharca asperula 5 0.4 
Bathyarca inaequisculpta 24 2.1 
Limopsis galathea 197 Tot 
Limopsis tenella 2 0.2 
Thyasira inflata 43 910 
Thyasira pygmaea 27 2.4 
Thyasira transversa 21 1.8 
Abra profundorum 2 0.2 
Kelliella atlantica 7 0.6 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 


(continued) 


Sample 


Sta. 196, 4,630 т, 10°29.0'S, 09°03.0’E 


Pristigloma nitens 
Deminucula atacellana 
Nuculoidea bushae 
Tindaria callistiformis 
Tindaria miniscula 
Ledella ultima 

Spinula hilleri 

Yoldiella fibula 
Yoldiella similes 
Portlandia abyssorum 
Silicula filatovae 
Bentharca asperula 
Bathyarca inaequisculpta 
Limopsis galathea 
Incerte cedis sp. 1 b 


Sta. Walda 0$03, 4,829 т, 20°03.8'S, 
07°59.9’E 

Limatula louiseae 

Limopsis galatheae 

Incerte cedis 


Sta. Walvis DS07, 5,157 m, 26°59.7’S, 
OIFOTMÉE 

Ledella ultima 

Yoldiella subcircularis 

Malletia abyssorum 

Dacrydium abyssorum 


Sta. Walvis DS09, 5,220 m, 26°59.9'S, 
01°06.7’E 

Ledella ultima 

Yoldiella subcircularis 

Limopsis galathea 

Kelliella atlantica 

Verticordia triangularis 

Incerte cedis 


Sta. Walvis DS08, 5225 m, 26°59.9’S, 
01078 E 

Ledella ultima 

Yoldiella subcircularis 


CAPE BASIN 


Эа. 161, 220m, 2253.0'5, 13°31.0°E 
Lucinoma filosa 


Sta. 186, 481 m, 22°57.0'S, 13°05.0’E 
Nucinella pretiosa 
Yoldiella hanna 
Bathyarca pectunculoides 
Limatula smithi 
Cyclopecten sp. a 
Thyasira alleni 


No. of 
Individuals 


O0 


NO 
N ND O1 © = = = NN ON = © OO N 


Wo — 
00 tr > 


ты 


Occurrence 
in % 


(continues) 


169 


170 


ALLEN 


(continued) 


Sample 


Thyasira carrozae 
Thyasira subovata minuta 
Thyasira transversa 
Thyasira sp. 186 
Mysella verrilli 

Mysella sp. 1 

Tellinidae sp. c 
Tellinidae sp. d 

Kelliella atlantica 
Kelliella elongata 
Thracia sp. 2 

Myonera tillamookensis 
Incerte cedis sp. 1 a 
Incerte cedis sp. 2 
Incerte cedis sp. 186 
Incerte cedis sp. 262 


Sta. 1168, 022 m, 23:00:05, 12°S80iE 


Nuculoma granulose 
Tindaria sp. 188 
Neilonella whoii 

Ledella acinula 

Ledella sandersi 

Yoldiella bilanta 

Yoldiella capensis 
Yoldiella hanna 
Portlandia minuta 

Silicula filatovae 
Bathyarca pectunculoides pellucida 
Limopsis cristata agg. 
Similipecten minor 
Thyasira alleni 

Thyasira carrozae 
Thyasira croulinensis 
Thyasira equalis 

Thyasira intermedius 
Thyasira subovata minuta 
Thyasira subovata subovata 
Thyasira succisa atlantica 
Thyasira tortuosa 
Thyasira transversa 
Thyasira sp. 28 

Thyasira sp. 188 a 
Thyasira sp. 188 b 
Galeommatoidea sp. 
Mysella ovata 

Mysella verrilli 

Tellinidae sp. b 

Tellinidae sp. d 

Kelliella atlantica 

Kelliella elongata 


No. of 
Individuals 


184 
133 


— 
EN 
O0 


© 
dl 


ass SS SS 


oO N 
sy (G0) ES) do, IN SS N N CO) 


Occurrence 


in % 


zer 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 1147 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Kelliella tenina 29 0.2 
Lyonsiella sp. 18 0.08 
Policordia densicostata 87 0.6 
Policordia sp. 5 0.03 
Laevicordia sp. 3 0.02 
Rhinoclama abrupta 340 22 
Luzonia simplex 224 1:8 
Myonera tillamookensis 5 0.03 
Protocuspidaria simplex 1 0.01 
Incerte cedis sp. 2 26 0.2 
Incerte cedis sp. 188 a 5 0.03 
Incerte cedis sp. 188 b 2 0.01 
Incerte cedis sp. 188 c 5 0:03 

Sta. 189, 1,014 т, 23°00.0’S, 12°45.0’E 
Neilonella salicensis 198 2.4 
Yoldiella capensis 918 1947 
Yoldiella curta 196 3.4 
Limopsis cristata lanceolata 2705 4.7 
Limatula smithi 55 0.9 
Cyclopecten sp. a 50 0.9 
Lucinoma filosa Y 0.1 
Thyasira alleni 250 4.3 
Thyasira carrozae 1810 34150 
Thyasira croulinensis 2 0.03 
Thyasira subovata subovata 40 0.7 
Thyasira tortuosa 20 0:3 
Thyasira transversa 362 6.2 
Mysella sp. 1 2 0.03 
Kelliella elongata 184 3,1 
Kelliella tenina 48 0.8 
Veneridae sp. h 8 0.1 
Policordia densicostata 1 0.02 
Halicardia flexuosa 1s 0.02 
Cuspidaria atlantica 20 0.4 
Rhinoclama abrupta 1 0.02 
Luzonia simplex 1 0.02 
Incerte cedis sp. 189 1464 2541 

Sta, 181, 1,659 m23°S, 12 31.TE 
Neilonella salicensis 52 2.5 
Ledella sandersi 14 07 
Yoldiella bilanta 1025 49.3 
Yoldiella curta 2 0.1 
Limopsis cristata lanceolata Z 0.1 
Limatula smithi 19 0.6 
Cyclopecten sp. a 20 1.0 
Lucinoma filosa 8 0.4 
Thyasira alleni 65 an 
Thyasira carrozae 174 8.4 
Thyasira croulinensis 48 23 
Thyasira eumyaria 1 0.05 


(continues) 


172 ALLEN 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Thyasira ferruginea 144 6.9 
Thyasira subovata subovata 29 1.4 
Thyasira succisa atlantica 30 125 
Thyasira tortuosa 40 1.9 
Thyasira transversa 68 312 
Leptonidae sp. w 36 dev 
Kelliella atlantica 2 0.1 
Veneridae sp.h 8 0.4 
Thracia sp. 3 8 0.4 
Policordia densicostata 2 0.1 
Policordia gemma 2 0.1 
Policordia insoleta 1 0.05 
Cuspidaria atlantica 10 0.5 
Luzonia simplex 240 11.6 
Incerte cedis sp. 262 34 ¡874 

Sta: 192, 2,154. 230205, 12 19.0°E 
Ledella sandersi 2 0.1 
Yoldiella bilanta 1697 81.2 
Yoldiella inconspicua africana 55 2:6 
Malletia johnsoni 3 0.1 
Bathypecten sp. e 20 1.0 
Lucinoma filosa 1 0.05 
Thyasira biscayensis 1 0.05 
Thyasira croulinensis 22 a 
Thyasira equalis 3 0.1 
Thyasira ferruginea 214 1055 
Thyasira subovata subovata 5 0.2 
Thyasira tortuosa 1 0.05 
Thyasira transversa at £3 
Veneridae sp. h 15 7 
Cuspidaria atlantica 1 0.05 
Luzonia simplex 19 0.9 

Sta. 194, 2,864 m, 22°54.0’S, 11°55.0’E 
Pristigloma nitens 3 0.4 
Phaseolus sp. a 14 2.0 
Ledella sandersi 1 0.1 
Yoldiella bilanta 2 0.3 
Yoldiella curta 2 0.3 
Yoldiella inconspicua africana Sl he 
Malletia johnsoni 2 O23 
Bentharca asperula 22 331 
Limopsis tenella 12 WA 
Bathypecten sp. e 10 1.4 
Thyasira brevis 99 13:9 
Thyasira ferruginea 452 659 
Thyasira succisa atlantica 29 ao 
Kelliella atlantica 2 2.9 


(continues) 


BIVALVIA OF THE DEEP ATLANTIC 173 


(continued) 
No. of Occurrence 
Sample Individuals in % 

Sta. Walvis DS05, 4,560 т, 33°20.5’S, 

02°34.9’E 
Ledella aberrata 2 MT 
Ledella ultima 34 28.8 
Portlandia abyssorum 1 0.9 
Yoldiella subcircularis 5 25 
Malletia abyssorum 9 7.6 
Limopsis galathea 2 ВА 
Dacrydium abyssorum 40 30 
Dacrydium sp. 2 HR 
Thyasira inflata 2 ats 
Thyasira sp. 2 ayer 
Kelliella atlantica 1 0.9 
Verticordia quadrata 1 0.9 
Myonera atlantica 18 1538 
Incerte cedis 1 0.9 

Sta. Walvis DS06, 4,585 m, 33°24.5’S, 

DIE 
Pristigloma alba 1 0.9 
Tindaria hessleri 1 0.9 
Ledella ultima 22 19.3 
Yoldiella jeffreysi 1 0.9 
Yoldiella subcircularis T 6.1 
Malletia abyssorum 183 115 
Bathyarca sp. 3 20 
Bentharca asperula 2 ТР 
Dacrydium abyssorum 43 ЗЕ 
Thyasira inflata 5 4.4 
Kelliella atlantica 1 0.9 
Myonera atlantica 3 2.6 
Incerte cedis sp. 1 1 0.9 
Incerte cedis sp. 2 1 0.9 

Sta. Walvis DS03, 4,657 m, 33°23.2’S, 

02°40.3’E 
Ledella ultima 1 100.0 

Sta. Walvis DS01, 5,240 m, 33°53.9’S, 

05°05.9’E 
Ledella ultima 2 33.3 
Limatula louiseae 1 16.7 
Kelliella atlantica 2 aoe 
Incerte cedis 1 16.7 

Sta. Walvis DSO2, 5,280 т, 33°54.7’S, 

OOF ae 
Ledella ultima 24 59.0 
Spinula hilleri 1 2.6 
Malletia abyssorum 9 Edi 
Malletia pallida 3 7.9 
Dacrydium abyssorum 2 6.5 


1 | o 
р Mit E 


MALACOLOGIA, 2008, 50(1-2): 175-218 


DWIGHT WILLARD TAYLOR (1932-2006): 
HIS LIFE AND MALACOLOGICAL RESEARCH 


Alan В. Kabat®* & Richard |. Johnson? 


ABSTRACT 


Dwight Willard Taylor (1932-2006) was a malacologist and paleontologist whose research 
on the systematics and biogeography of freshwater gastropods, particularly the Hydrobiidae 
and Physidae, resulted in numerous taxonomic innovations and led to extensive research 
by others. His biogeographical analyses of the distribution of freshwater mollusks, particularly 
from western North America, were provocative and stimulating. His research on endangered 
or threatened species was influential in the conservation of those species and their habitats. 
He published 65 papers and 9 abstracts, and authored numerous internal reports to federal 
and state agencies, primarily for the U.S. Geological Survey and the U.S. Fish and Wildlife 
Service. He described 132 taxa, comprising 12 family-level taxa, 31 genus-level taxa, and 
89 species; most of his taxa were in the Hydrobiidae (54) and Physidae (39), and he was 
involved in the founding and early development of Malacologia. 

Key words: biogeography, Hydrobiidae, Physidae, Dwight W. Taylor. 


BIOGRAPHY AND ANALYSIS OF 
TAYLOR'S RESEARCH 


Dwight Willard Taylor’s malacological re- 
search on freshwater mollusks led to a number 
of significant accomplishments. This biographi- 
cal essay discusses his life, his professional 
career, and his research. 

He was born on January 18, 1932, in Pasa- 
dena, a suburb of Los Angeles, California. His 
parents were Daniel Dwight Taylor (died 1969), 
an engineer at Beckman Instruments, and Sa- 
rah Willard Taylor (died 2002). Through his 
mother, the family was wealthy twice over. His 
maternal grandfather was Henry K. Willard of 
Washington, D.C., whose father was the founder 
of the Willard Hotel, the well-known luxury hotel 
on Pennsylvania Avenue, midway between the 
White House and the Smithsonian’s National 
Museum of Natural History (Wallace & Carr, 
1986). His maternal grandmother, Helen Willard, 
was the daughter of E. Southard Parker, the 
president of several banks in Washington, D.C. 
(Anonymous, 1966a, b). He was raised in Altade- 
na, near Pasadena, in a baronial Spanish-Medi- 
terranean mansion, at 1955 Mendocino Lane, 
in the foothills of the San Gabriel Mountains. 


High School Years (1945-1949) 


Taylor was educated at the Webb School, a 
private boarding school in Claremont, Califor- 
nia, in the foothills east of Pasadena. The 
Webb School, then as now, is unique among 
high schools in having a significant focus on 
paleontology, which arose from the interests 
of a charismatic biology teacher, Raymond 
Manfred Alf (1903-1999). Mr. Alf, during a 
school field, trip, was the first to discover the 
fossil of a peccary, or wild pig (Jameson & 
McMillin, 1985; Lofgren, 2000, 2005; Woo, 
1999). Mr. Alf was a mentor to several gen- 
erations of students, a number of whom went 
on to distinguished careers in paleontology, 
notably including Malcolm Carnegie McKenna 
(class of 1948), longtime curator of vertebrate 
paleontology at the American Museum of Natu- 
ral History, and Taylor (class of 1949)'. Taylor 
was later the best man at McKenna’s wedding. 

In 1968, the Webb School opened the Alf 
Museum of Paleontology to house the sizable 
collections amassed by Mr. Alf and his stu- 
dents; this remains the only high school mu- 
seum in the U.S.A. that is formally accredited 
(www.alfmuseum.org). The funds for building 


“Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. 
5124 Chestnut Hill Road, Chestnut Hill, Massachusetts 02467-1310, U.S.A. 


“Corresponding author: alankabat@aol.com 


‘In the 1980s, Taylor received significant research funding from the Philip McKenna Foundation, which was established 
by the uncle of Malcolm McKenna, and derived its endowment from Pittsburgh steel money. 


176 KABAT & JOHNSON 


the Alf Museum came primarily from Webb 
School trustee George Getty’. Taylor’s parents 
donated funds for the “Dwight Willard Taylor 
Laboratory of Biological Sciences,” a teach- 
ing laboratory and classroom adjacent to the 
museum exhibits. He honored Mr. Alf by de- 
scribing Helminthoglypta alfi Taylor, 1954, a 
fossil terrestrial gastropod; the type material 
included specimens collected by Mr. Alf and 
his students. Don Lofgren, the Director of the 
Alf Museum, aptly described Mr. Alf’s enthusi- 
asm for teaching, based on the recollections 
of former students: 

“Ray employed many innovative teaching 
methods that he had developed over the years 
for his students or The Boys; he always re- 
ferred to his former students as The Boys 
whether they were from the class of 1939 or 
1969 ... Ken De Nault ‘61 reflected on another 
of Ray’s simple but effective teaching meth- 
ods ... ‘Finally, when | did get it right, | received 
the highest praise a Peccary Man could re- 
ceive — Good Boy. Through all of this Ray was 
teaching. The lesson was that what was in our 
pan was what we created. The only one re- 
sponsible was our self. This was an important 
lesson for life ahead of us.’ Ray led by ex- 
ample. This did not escape the notice of his 
students ... ‘His exhortations to have the fight- 
ing heart and to know that each day was a 
test of a man’s character were far from empty 
rhetoric’.” (Lofgren, 2000: 2; italics in original). 

During his high school years, Taylor also 
came to know S. Stillman Berry (1887-1984), 
a malacologist who published extensively on 
nearly all the classes and orders of mollusks 
and edited the Leaflets in Malacology from his 
home research laboratory in Redlands, south- 
east of Pasadena, and John О. Burch (1894— 
1974), ashell and book dealer in Los Angeles, 
who edited the Minutes of the Conchological 
Club of Southern California. 

While at the Webb School, Taylor began to 
assemble what became a sizable working li- 
brary of malacological literature, with an em- 
phasis on publications treating non-marine 
mollusks. Taylor was among the earliest cus- 
tomers of one of us (Johnson), who as a col- 
lege student had started selling mollusk books 
from his parents’ home. 


The Mollusks of Nantucket 


While growing up, the Taylor family, includ- 
ing Dwight and his two sisters (Sarah and 
Margaret), spent their summers at Grandma 
Willard’s house on Nantucket Island, Massa- 
chusetts, where she maintained a summer 
residence from the 1920s through her death 
in 1966. Nantucket is unusual among summer 
resorts in having a historically important sci- 
entific research station, the Nantucket Maria 
Mitchell Association, named after the first fe- 
male astronomer in the U.S.A., and which has 
a children’s nature program. Taylor explored 
the diverse molluscan fauna of Nantucket, both 
marine and terrestrial, starting under the dot- 
ing tutelage of Miss Grace Wyatt, a summer 
scholar at the Maria Mitchell Association who 
ran the summer programs, which included 
extensive nature walks (Anonymous, 1955; 
Drake, 1968). 

Taylor also began corresponding with Will- 
ат J. Clench (1897-1984), then curator of 
mollusks at the Museum of Comparative Zo- 
ology, and one of us (Johnson), then an un- 
dergraduate at Harvard. His letters, written 
from both Nantucket and the Webb School, 
reveal him to be enthusiastic about collecting 
mollusks, and remarkably interested in their 
taxonomy and biogeography. For example, in 
1948, at the age of 16, he asked Clench how 
to differentiate the nominal species of the 
marine bivalves Astarte and Gemma, and said 
that he had “started working on Amphimelania, 
more for practice than anything else, and am 
dismayed at the chaos | have found,” largely 
because Bourguignat had described so many 
species, but he concluded that “it is an awful 
lot of fun, and | heard of people | had never 
thought of before (Spiridion Brusina, for ex- 
ample).” (D. W. Taylor to W. J. Clench, in litt., 
July 31, 1948). 

In 1949, at the age of 17, while a senior at 
the Webb School, he reached the national fi- 
nals of the Westinghouse Science Scholarship 
Program. This is a prestigious talent competi- 
tion that (in 1949) culminated with 40 bright 
high school students from across the country, 
selected from a much larger pool of applicants, 
traveling to Washington, D.C. to present the 


“George Franklin Getty (1925-1973) was the only one of the four sons of oilman J. Paul Getty (1892-1976) to attend the 
Webb School (class of 1942), and presumably his exposure to geology under Mr. Alf helped him serve as Executive Vice 
President of Getty Oil Company. George died young, shortly after helping start the Alf Museum (Pearson, 1995; West, 


1973). 


DWIGHT WILLARD TAYLOR 17% 


CE NEWS LETTER 


FIG. 1. Cover, Science News Letter, showing Taylor as one of the two top finalists in the Ninth Annual 
Science Talent Search for Westinghouse Science Scholarships in March 1949 (Anonymous 1949d). 


178 KABAT & JOHNSON 


FIG. 2. The Ninth Annual Science Talent Search 
for Westinghouse Science Scholarships in March 
1949. Taylor is standing in front of his hand-drawn 
map of Nantucket, with mollusk specimens on 
the table and a jar of specimens in his left hand 
(Taylor 1949b, courtesy of Paul Valentich-Scott). 


results of their research projects, and to meet 
with President Truman in his office at the White 
House, members of Congress, and govern- 
ment researchers. This competition continues 
today, now sponsored by the Intel Corporation. 

From March 3 to 7, 1949, Taylor was in 
Washington, D.C., along with 39 other high 
school students, who were selected from a 
pool of 16,218 applicants (Anonymous, 1948, 
1949а-е). His project was titled “A malaco- 
logical survey of Nantucket island, Massachu- 
setts,” and his extensive collecting increased 
the number of known molluscan species from 
45 to 120. The brochure of this competition 
included an excerpt from his essay as one of 
only two essays selected for publication (Tay- 
lor, 1949b). This essay reveals a precocious 
scholar who used his extensive collections to 
draw inferences about the ecological and bio- 
geographical relations of the marine mollusks 


of Nantucket. Taylor also discovered two new 
exposures of fossil-bearing strata on the is- 
land, aided in that search by the paleontologi- 
cal skills he learned from Ray Alf. 

The press coverage of this competition was 
quite adulatory, particularly once Taylor was 
selected as the recipient of the Grand Scholar- 
ship of $2,800 (the remaining competitors re- 
ceived smaller scholarships ranging from 
$2,000 to $100). For example, the Science 
News Letter described Taylor as “the nation’s 
top young scientist of 1949" and featured him 
on the cover (Anonymous, 1949d) (Figs. 1, 2), 
while Time magazine described him as “top of 
the crop” (Anonymous, 1949e). He was not 
modest about his accomplishments: “To Dwight, 
the announcement came as no great surprise. 
... he has also been studying the distribution, 
taxonomic position and ecology of mollusks in 
Southern California. Where would all this lead 
him (after four years at the University of Michi- 
gan)? ‘Oh, ГИ probably end up in some univer- 
sity museum or something. One can’t live on 
just nothing’.” (Anonymous, 1949e). 

The Webb School was justifiably proud of 
this accomplishment, since he was their first 
student to be a Westinghouse finalist, let alone 
the national champion. The school’s yearbook 
for 1949 includes two photos of the “parade” 
upon his return from Washington, D.C., in 
which he rode in the back of an open convert- 
ible through the campus, underneath a ban- 
ner, “Welcome Home Dwight” and surrounded 
by cheering students, much as in a parade for 
the astronauts returning from the moon (Figs. 
3, 4). In 1954 and 1955, two other Webb stu- 
dents, Patrick Muffler and David Fleishhacker,? 
also became Westinghouse finalists for their 
research on fossil mammals (oreodonts), and 
Mr. Alf arranged to have all three (Taylor was 
then in graduate school at Berkeley) photo- 
graphed together on campus, and the poster- 
sized photograph is on display at the Alf 
Museum (Fig. 6). In retrospect, this excessive 
adulation at an early age may have affected 
Taylor’s perception of his own abilities in com- 
parison to those of his colleagues. 

He concluded his Westinghouse essay by 
noting that he planned to investigate those 


®Leroy John Patrick Muffler (class of 1954) obtained his Ph.D. in geology at Princeton University in 1962, and spent his 
career at the U.S. Geological Survey, where he was promoted to Western Regional Geologist, and later Acting Director 
of the Menlo Park office. David Fleishhacker (class of 1955) became a teacher and administrator at several private schools 
in California. He was a scion of the well-known San Francisco family; Herbert Fleishhacker (1872-1957), a banker, founded 
the Fleishhacker Zoo, now the San Francisco Zoo, and built Fleishhacker Pool, then the world’s largest swimming pool 


and now the parking lot for the zoo. 


DWIGHT WILLARD TAYLOR 109 


FIGS. 3, 4. At the Webb School (Claremont, California) in March 1949. 
FIG. 3 (Top): The “parade” at the Webb School upon Taylor’s return from 
the Westinghouse competition. Photo from E/ Espejo (yearbook) for 1949 
(courtesy of Don Lofgren); FIG. 4. (Bottom): Thompson Webb, Director of 
the Webb School, presenting Taylor with the “key” to the Webb School. 
Photo from E/ Espejo (yearbook) for 1949 (courtesy of Don Lofgren). 


fossil outcrops in greater detail, “eventually 
undertaking a comparison of the Recent and 
Pleistocene faunas’, which “will lead to a study 
of Quaternary geologic and climatic changes 
in the area” (Taylor, 1949b: 20), but did not 
publish that study. Of the remaining competi- 
tors in 1949, the only other recognizable name 
is Walter Gilbert, who went on to become chair- 
man of the Cellular & Developmental Biology 
department at Harvard, and who patented his 


Nobel-prize winning method for sequencing 
DNA. 

Taylor graduated from the Webb School in 
1949. The school’s yearbook, E/ Espejo, in- 
cluded a half-page for each graduating senior, 
consisting of a photograph and a twelve-line 
rhyming poem about that student. The year- 
book photograph shows him examining an ar- 
ticle with illustrations of snail shells (Fig. 5), 
and the accompanying poem is worth quoting: 


180 KABAT & JOHNSON 


There also came a lad by name Dwight Taylor, 
Who from his interests now is known as “Snailer.” 
The wisdom of his words is not denied, 

Though sometimes they are used to cut or chide. 
Oft El Espejo, Blue and Gold receive 

His toil; in his lush garden we perceive 

The greenest lawn for leagues — or so he states. 
Although Dwight’s grades are not the top, he rates 
As one whom Westinghouse gave passing fame 
And who from far and wide received acclaim. 

On deserts wide for fossils he does search; 

And near both stream and pond pursues research. 


Pete Akin, one of Taylor’s classmates, told 
one of us that Taylor, who was on the year- 
book staff, not only wrote his own poem, but 
also wrote most of Akin’s poem. Taylor’s poem 
recognizes his passion for fossils, freshwater 
snails, and writing for a public audience. 


Undergraduate Years (1949-1953) 


In his senior year, Taylor was faced with the 
choice of where to apply for college. He could 
have followed Malcolm McKenna to the Uni- 
versity of California, Berkeley. McKenna, al- 
though only an undergraduate, had already 
learned so much paleontology from Mr. Alf, that 
the faculty made him the teaching assistant for 
the graduate courses in paleontology (Bell, 
2004: 11). Alternatively, Taylor could have ap- 
plied to Harvard University, since he had used 
the Museum of Comparative Zoology mollusk 
collections and library for identifying his Nan- 
tucket specimens, ably guided by William 
Clench and Ruth Turner, or he could have ap- 
plied to Stanford University, the alma mater of 
Stillman Berry, and where Myra Keen was 
mentoring several generations of malacolo- 
gists. However, Taylor chose to do his under- 
graduate studies at the University of Michigan. 

Although Michigan has extensive non-marine 
mollusk collections, then curated by Henry van 
der Schalie (1907-1986), his interest in Michi- 
gan was also motivated by the opportunity to 
work with Claude William Hibbard (1905- 
1973), the vertebrate paleontologist whose 
novel methods for collecting small-sized mam- 
malian fossils revolutionized mammalian pa- 
leontology by greatly expanding the scope of 
recoverable specimens (Adler, 2007: 191- 
192). Hibbard was affectionately known as 
“Hibbie” to his students, who regarded him as 
a “larger-than-life character,” and his “inspired 


(Webb School, 1949: 26) 


teaching and infectious enthusiasm for the pre- 
historic world” motivated even the Zoology stu- 
dents to pursue paleontological research. (K. 
К. Adler to A. R. Kabat, in litt., Jan. 5, 2008). 
Taylor’s studies and field work under Hibbard 
allowed him to become extensively familiar with 
the stratigraphy and geologic history of the 
Midwest states, including Kansas, Nebraska, 
and Oklahoma, and westwards to Wyoming 
and Montana (Anonymous, 1957). He also 
enjoyed interacting with Harold (Hal) Harry 
(1921-1995), a graduate student who became 
“a willing advisor, stimulating field trip compan- 
ion, and generous host” (Taylor, 2002b: 159). 


FIG. 5. Taylor examining article with illustrations 
of gastropod shells, Webb School yearbook pho- 
tograph, 1949 (courtesy of Don Lofgren). 


DWIGHT WILLARD TAYLOR 181 


FIG. 6. The Webb School's finalists in the Westinghouse Science Talent Search of America (left to 
right): Patrick Muffler (class of 1954); David Fleishhacker (class of 1955); Dwight W. Taylor (class of 
1949). This photograph was probably taken in 1955, and a poster-sized version is on display at the 


Alf Museum (courtesy of Don Lofgren). 


While an undergraduate, Taylor published 
two short papers in Leaflets in Malacology, 
edited by Stillman Berry. The first described 
three new species of Pyrgulopsis from the 
Colorado Desert region of southern California 
(Taylor, 1950), and the second discussed the 
freshwater mollusks of Yellowstone National 
Park, Wyoming (Taylor, 1952). He candidly 
admitted, 20 years later, that two of the new 
species in his 1950 paper were junior syn- 
onyms, but he gratefully recognized Berry’s 
encouragement: “With pleasure | dedicate this 
work to Stillman Berry, a long-time friend and 
mentor. Perhaps it will make up in part for the 
time and trouble he took in encouraging me, 
from high-school days on ... Like many bud- 
ding zoologists, | thought in my early days that 
finding a new species and publishing a paper 
were about the ultimate achievements in life. 
Well, you have to get it out of your system. 
And Stillman helped me ... Although | never 
told him so, | realized later that this was a pretty 
half-baked job. Two of the three species | 


thought | was naming are synonyms, and be- 
long to a different genus, and probably are the 
same species anyway. The illustrations are 
rather sketchy. And one should not describe 
Hydrobiidae from shell alone until a fairly thor- 
ough search for living snails has been com- 
pleted. But | like to think that Stillman saw | 
could go beyond species-naming, and that the 
sooner | passed through that phase the bet- 
ter.” (Taylor, 1970). 

In 1951, while an undergraduate, Taylor ob- 
tained a copy of Alfred Russell Wallace’s 1876 
treatise: “The Geographical Distribution of 
Animals.” He was proud to have this impor- 
tant book, and inscribed in his copy: “| have 
wanted this work for a long time. | am now 
even more sure that | shall be a zoographer. If 
| may succeed with my data as Wallace did 
with his, | shall be content.” (Barrientos & 
Springer, 2007: ix). 

Taylor withdrew from Michigan, ostensibly 
because he did not like the cold winters, and 
transferred to Pomona College in southern 


182 KABAT & JOHNSON 


California for his sophomore and junior years. 
However, he then returned to Ann Arbor to 
complete his undergraduate studies. During 
his last year, he overlapped with George Alan 
Solem (1931-1990), who began his graduate 
studies at Michigan in 1952, and devoted his 
career to the study of terrestrial pulmonates, 
but had a falling out with Taylor. 

It was around this time that Taylor’s parents 
expressed their strong expectations that he 
would use his academic abilities to become a 
physician or a lawyer, and not devote his life 
to Cenozoic malacology. As he later told 
Hibbard, in the ensuing arguments with his 
parents, “he dissociated himself from the fam- 
ily, but was given one million dollars with the 
understanding that there would be no addi- 
tional inheritance and no family contact” (G. 
К. Smith to A. R. Kabat, in litt., June 19, 2007). 


Graduate Studies (1953-1957) and Work at 
the Survey (1955-1967) 


In 1953, Taylor earned his B.S. degree from 
the University of Michigan, and he promptly 
enrolled in the graduate program at the Uni- 
versity of California, Berkeley, where he re- 
ceived his Master’s degree in 1954 and his 
Ph.D. degree in 1957. One of his roommates 
at Berkeley was William K. Emerson, later 
curator of mollusks at the American Museum 
of Natural History. The Master’s thesis, on six 
Pliocene and Pleistocene mollusk faunas from 
Kansas and Nebraska, was an outgrowth of 
his undergraduate collecting work with 
Hibbard. 

While still in graduate school, he published 
on the non-marine mollusks from the Upper 
Miocene Barstow Formation, based primarily 
on specimens obtained during the Webb 
School field trips with Mr. Alf and his fellow 
students. In addition to describing Helmintho- 
glypta alfi, named for his mentor, he also de- 
scribed the oldest fossil slug known from North 
America, Craterarion pachyostracon Taylor, 
1954 (Arionidae). It may come as a surprise 
that slugs even have a fossil record (Tracey 
et al., 1993: 163), but this species is moder- 
ately common in the Barstow deposits. 

In 1955, Taylor joined the staff of the U.S. 
Geological Survey (USGS), Paleontology & 
Stratigraphy Branch, then located at the U.S. 
National Museum (now the National Museum 
of Natural History), Smithsonian Institution, 
Washington, D.C. The Paleontology & Stratig- 
raphy Branch was then the world’s largest 
paleontological research program, with sev- 


eral dozen paleontologists and numerous tech- 
nicians and assistants (Dutro, 2004). When he 
was at the Survey, J. Thomas Dutro (an ex- 
pert on brachiopods) was the Branch Chief of 
the Paleontology & Stratigraphy Branch, and 
the other Cenozoic molluscan paleontologists, 
all primarily interested in marine mollusks, in- 
cluded Wendell Woodring (1891-1983) and 
Harry Ladd (1899-1932), both then approach- 
ing retirement, and Druid Wilson (1906-2002). 

Even before joining the Survey, Taylor pub- 
lished in its Professional Paper series (Taylor 
1954), and amassed an impressive publica- 
tion record while at the Survey. Taylor returned 
to Berkeley in January 1957, where he suc- 
cessfully defended his Ph.D. thesis, a lengthy 
analysis of the Late Cenozoic paleoecology 
and molluscan faunas of the High Plains (Tay- 
lor, 1957; published as Taylor, 1960; Hibbard 
& Taylor, 1960). 

While at the Survey, Taylor prepared numer- 
ous internal agency reports, known as “Exami- 
nation & Report,” which identified fossils that 
were collected by others, primarily in the west- 
ern United States. The originals of these type- 
script reports were given to the sender of the 
specimens, with a carbon copy retained in the 
USGS archives. His reports remain valuable 
in their detailed discussion of the ecological 
habitat, biogeography, and stratigraphy of the 
species, particularly as many of his observa- 
tions were never formally published. Some of 
Taylor's reports have been cited in the subse- 
quent literature, notably by Malde & Powers 
(1962) in their studies of the Snake River Plain 
stratigraphy. These USGS reports are cata- 
loged in a separate annotated bibliography 
(Kabat, in prep.). 

In 1958, Taylor wrote that he and most other 
Survey paleontologists would be unable to do 
field work, because “a low budget and the ter- 
mination of support by the AEC [Atomic En- 
ergy Commission] and Army have squeezed 
us rather badly. I’ll be going on leave and tak- 
ing Russian in summer school ... Ability to read 
Russian will also be quite useful, and I’m glad 
| can get started learning it. Don’t misunder- 
stand; Га much rather be in the field.” (D. W. 
Taylor to W. К. Emerson, in litt., May 29, 1958). 
In later years, Taylor's knowledge of Russian 
allowed him access to the extensive Russian 
literature on the trans-Beringian distribution of 
freshwater mollusks, a subject of great bio- 
geographic interest. 

Working at the Survey was his first introduc- 
tion to the regular workplace world, where he 
had to interact on a daily basis with supervi- 


DWIGHT WILLARD TAYLOR 183 


sors, colleagues, and subordinates. Hitherto, 
he had cloistered himself in an academic en- 
vironment, where he only had to answer to his 
faculty advisors, who may have been eccen- 
tric or otherwise able to tolerate his personal- 
ity. Now, however, he was placed right in the 
middle of a sizable government bureaucracy, 
with layers of administrators, and mountains 
of paperwork. 

Although Taylor remained respectful of se- 
nior mentors at the Survey, such as Ladd and 
Woodring, he was barely able to tolerate his 
supervisor, Dutro, who was only a decade 
older, and his relationships with some of the 
other geologists and paleontologists rapidly 
deteriorated. He was seemingly unable to have 
a professional discussion about stratigraphical 
issues that did not end up in a shouting match 
that could be heard down the hallway. Taylor 
could seldom admit that his colleagues’ opin- 
ions might be valid. 

Among the requirements of working at the 
Survey were that all manuscripts had to be 
submitted for internal review and approval 
before they could be submitted for publication. 
Taylor chafed under these restrictions, which 
he viewed as delaying his output, and did not 
want to have to respond to suggestions for 
revisions to his manuscripts. Thus, Taylor took 
advantage of the fact that he was also a Re- 
search Associate at the Museum of Zoology, 
University of Michigan, and several of his pa- 
pers published during his years at the Survey 
make no mention of the Survey, let alone pub- 
lication approval, instead listing his only affili- 
ation as Michigan. Interestingly, the Survey’s 
requirements for internal review are still con- 
troversial to some scientists who view this as 
censorship, as recently reported in the Wash- 
ington Post (Eilperin, 2006). 

Taylor needlessly antagonized his col- 
leagues to the point where some repeatedly 
complained to Dutro (who, of course, already 
knew of this issue), yet Dutro could not solve 
this problem. Finally, Dutro asked Ladd how 
to deal with him, and after much discussion, 
Ladd chuckled and perceptively said to the 
effect that, “You can’t do anything with Dwight, 
as he is an idiot savant,” meaning that he was 
brilliant about mollusks, but unable to work with 
others. 


The Smithsonian Mollusk Department 
During Taylor’s tenure at the Survey, the 


Mollusk Department researchers included 
Harald Rehder (1907-1996) and Joseph 


Morrison (1906-1983) for that entire time, Paul 
Bartsch (1871-1960) having retired in 1946 
and К. Tucker Abbott (1919-1995) having 
moved to Philadelphia in 1954. Later came 
Joseph Rosewater (1928-1985) from 1960 
onwards, and Kenneth Boss of the U.S. Fish 
& Wildlife Service from 1963 to 1966. Morrison, 
who had published on brackish-water 
hydrobiids (Kabat & Hershler, 1993: 75), and 
who was an inveterate collector of mollusks, 
had some influence on Taylor, at least in the 
earlier years. 

However, Taylor found the Mollusk Depart- 
ment, before the arrival of Rosewater and Boss, 
to be moribund, writing in 1958 that the Uni- 
versity of Michigan was much more active, 
based on a recent visit: “It was a reassuring 
and stimulating experience to see an institu- 
tion where research on Recent mollusks is 
being carried on. Association with the National 
Museum tends to get one down gradually.” (D. 
W. Taylor to W. K. Emerson, in litt., May 29, 
1958). In 1963, the new East Wing of the U.S. 
National Museum opened, with more than half 
of the space devoted to paleontology, includ- 
ing the extensive Survey collections. The Ceno- 
zoic mollusk collections and staff moved to the 
5" floor, which they shared with the Recent 
mollusk collections and staff. The Recent mol- 
lusk collections took up most of the eastern 
half of the 5" floor, with the Cenozoic type 
specimens at the northern end of that space, 
a situation that remained unchanged until 2000, 
when the Recent mollusk collections and staff 
moved to the West Wing to be with the rest of 
the Department of Invertebrate Zoology. 

His professional relations with his colleagues 
at the Smithsonian who worked on Recent 
mollusks were more amicable than with the 
Survey paleontologists, probably since he did 
not have to discuss stratigraphic matters with 
them. Yet, Taylor seldom joined those col- 
leagues or their visitors for their regular 
lunches in the Mollusk workroom, at which all 
matters malacological and otherwise were rou- 
tinely discussed. 

Among the numerous visitors to the Recent 
mollusk collections during Taylor’s years at the 
Survey was Bengt Hubendick (born 1916), the 
Swedish malacologist who had published an 
exhaustive monograph on the freshwater pul- 
monate family Lymnaeidae. Hubendick syn- 
onymized nearly all of the nominal genera 
under Lymnaea and synonymized hundreds 
of the described species (Hubendick, 1951). 
In advance of Hubendick’s visit, Boss walked 
down the hall to arrange for Hubendick to meet 


184 KABAT & JOHNSON 


with Taylor, given their mutual interests in 
freshwater mollusks and that Taylor had re- 
cently published on the Lymnaeidae (Taylor 
et al., 1963). However, as soon as Boss said 
that Hubendick was coming, Taylor made furi- 
ous statements about Hubendick’s research, 
and told Boss, in no uncertain terms, that “if 
you bring that SOB into my office, | will throw 
him out!” Thus, when Hubendick arrived, Boss 
had to arrange a delicate choreography to 
ensure that Hubendick never even walked by 
Taylor’s office, and always took the “long way 
around” to get to the mollusk collection from 
the elevators, all without telling Hubendick 
why. Four decades later, in what was to be 
the last paper before his death, he remarked 
on the “pretentious work by Hubendick” (Tay- 
lor, 2004а: 279). 


Life in Washington, D.C. 


When Taylor first moved to Washington, he 
lived with Grandma Willard, who resided in one 
of the largest mansions in the prestigious 
Kalorama Circle neighborhood, near Rock 
Creek Park. Her residence, at 2425 Wyoming 
Avenue, N.W., exceeds 10,000 square ft on 
three floors, and overshadows its somewhat 
smaller neighbor, which is now the residence 
of the Austrian Ambassador. 

According to the city directories, Taylor later 
bought a townhouse on Capitol Hill, at 525 — 
4" Street, S.E., several blocks from the Library 
of Congress. As Boss lived in a much more 
modest apartment nearby, Taylor gave him a 
grand tour of the townhouse. Boss, after ad- 
типо its features, jokingly said, “Well, if | save 
my pennies, | hope to be able to buy a place 
like this.” Taylor’s immediate rejoinder, “Oh, 
you'll never be rich!” 

While living in D.C., Taylor became a mem- 
ber of a private gentlemen’s key club, the 
Gaslight Club (1020 — 16" Avenue, N.W.), 
founded in Chicago, and with branches in New 
York and D.C. He proudly took as guests those 
few of his colleagues at work whom he toler- 
ated (eventually, only Boss met that criterion), 
along with several out-of-town visitors, prima- 
rily Arthur Merrill and Richard Petit. Boss and 
Petit well remember the details of the Gas- 
light Club. One entered an ordinary-looking 
building, and went into the men’s bathroom, 
located off the lobby. At the back was a wall 


on which the member would knock in a coded 
pattern. A hidden panel in the wall would open, 
allowing the guard on the other side to see 
who was in the bathroom, as in a Prohibition- 
era “speakeasy.” Upon being satisfied that the 
person was a member of the club, the guard 
would open a secret door, allowing Taylor and 
his guests to go upstairs to the gentlemen's 
club, on the second floor. 

The exclusive Gaslight Clubs later inspired 
the Playboy clubs that became popular in the 
1960s. There was a bar, a dance floor with a 
live band, and tables with comfortable leather 
seats. The men’s room within the club was 
noted for its large copper-lined urinal that ran 
along an entire wall. The urinal was filled with 
ice cubes, and had a bank of telephones on 
the wall above the urinal, so that the inebri- 
ated customers could, and did, make tele- 
phone calls while relieving themselves. The 
generously endowed and scantily attired host- 
esses were numerous and highly attentive, 
selling overpriced cocktails. One hostess, in 
particular, always served Taylor, who proudly 
told Merrill, “She’s in love with me.” Merrill, 
older and wiser in such matters, responded, 
to no avail, “Dwight, she’s in love with your 
money!” This did not stop Taylor from vastly 
over tipping the hostess who, fortunately, 
never married him. 


Founding of Malacologia 


In 1959, John Burch at Michigan proposed 
a new malacological serial, to be called 
Malacologia, to provide a venue for lengthy 
research papers, including those with detailed 
anatomical or stratigraphical analyses, given 
that the other malacological serials then pub- 
lished in the USA either had only short papers 
(The Nautilus) or were narrowly focused on 
systematic monographs (Johnsonia). Taylor 
later claimed, incorrectly, that he came up with 
the idea for this journal, based on conversa- 
tions he had with Burch at a disreputable bar 
on Capitol Hill at 8" Street S.E., located near 
the U.S. Marine Corps barracks, and known 
for its regular fights among the customers. 
Burch instead dated the formal founding of 
Malacologia to a discussion in June 1961 held 
by Burch, Melbourne Carriker, Robert 
Robertson and Taylor (Burch & Huber, 1968: 
29). 


4As this paper was published in a bilingual journal with English and Spanish texts in parallel columns, this phrase was 
carefully translated as “del pretencioso trabajo de Hubendick.” 


DWIGHT WILLARD TAYLOR 185 


At the 1961 meeting of the American Mala- 
cological Union, Burch, Taylor, and several 
other malacologists formed the Institute of 
Malacology (the name was Taylor’s idea), the 
sponsor for Malacologia (Carriker came up with 
the journal’s name). Burch prepared a grant 
application, which the National Science Foun- 
dation approved, covering journal expenses for 
the first three years. The first issue of this new 
journal, published in November 1962, led off 
with a carefully annotated and highly useful “An 
outline of gastropod classification” (Taylor & 
Sohl, 1962) co-authored by Taylor and Norman 
Sohl, a Survey paleontologist who worked on 
Mesozoic gastropods. In that issue, Taylor was 
listed as the President of the Institute of Mala- 
cology. 

However, his involvement with the Institute 
of Malacology took a downward turn. While 
Burch was away on travel, he entrusted Taylor 
with the editing and publishing of volume 3 of 
Malacologia. He used that opportunity to pub- 
lish a paper that described a new genus and 
species, Fontelicella californiensis, a minute 
freshwater gastropod (Hydrobiidae), based on 
a single type specimen, but with no illustration 
of the type specimen (Gregg & Taylor, 1965). 
Burch, upon his return, was concerned that this 
paper was published, since it would have ben- 
efited from careful review and illustrations of 
the new species. Also, another author’s manu- 
script was “bumped” in order to publish this 
paper. Boss tried to act as an intermediary, but 
was told by Taylor that, “my descriptions are 
so perfect that no illustrations are needed.” 
Taylor later resigned from the Institute and 
ceased any further involvement with 
Malacologia. 

Wendell Gregg (1898-1979) was a physician 
who had an extensive collection of the non- 
marine mollusks of southern California and 
adjacent regions. After Gregg and Taylor pub- 
lished their 1965 paper, they prepared a lengthy 
manuscript on other species referable to 
Fontelicella but, as Walter Miller (1918-2000) 
of the University of Arizona later wrote, “Unfor- 
tunately, he [Gregg] and Dwight had a bitter 
disagreement on how to proceed after their first 
publication and Gregg refused any further col- 
laboration” (W. Miller to R. Hershler, in /itt., Mar. 
3, 1988). 


Professional Societies 
Although Taylor did not attend American 


malacological meetings in his later years, in 
the 1950s and 1960s he did attend several 


meetings of the American Malacological Union 
(AMU, now the American Malacological Soci- 
ety), and its offshoot, the AMU-Pacific Division 
(today’s Western Society of Malacologists, 
WSM). Specifically, he joined the AMU in 1946, 
and his attendance was recorded at the AMU 
meetings in 1947 (Pacific Grove), 1950 (Chi- 
cago), 1958 (Ann Arbor), 1966 (Chapel Hill), 
and 1967 (Ottawa); and the AMU-Pacific Divi- 
sion in 1966 (Seattle) and 1967 (Pacific 
Grove). He, or his co-authors, presented pa- 
pers at the 1966 and 1967 meetings of the 
AMU-Pacific Division, and the 1969 meeting 
of the WSM. Taylor submitted a paper on en- 
dangered Western freshwater mollusks to the 
symposium on “Rare and Endangered Mol- 
lusks of North America” at the 1969 AMU meet- 
ing, but did not present his paper at that 
meeting. Clif Coney subsequently added 
Taylor’s name to an abstract for the 1985 WSM 
meeting, after Coney had presented a paper 
under his name alone on freshwater bivalves. 
In 1962, Taylor attended the first European Ma- 
lacological Congress in London, now known 
as the Unitas Malacologica (Fig. 7). 

In 1967, Taylor served as Second Vice Chair- 
man of the AMU-Pacific Division. In that capac- 
ity, he informed several colleagues that he was 
going to organize and edit a book, Malacology 
in Western America, which was to include bib- 
liographic chapters on various topics, as well 
as a world-wide bibliography of published type 
catalogs. This collaborative project did not pro- 
ceed further, but did result in two bibliographies 
authored by Taylor alone (Taylor, 1970b, 1975). 
Ordinarily, Taylor’s position as Second Vice 
Chairman would have led to his becoming 
Chairman, and hosting the annual meeting, but 
he was not nominated to an officer position in 
1968, and instead ceased his involvement with 
that organization. Taylor was present at a meet- 
ing on December 10, 1967, that led to the for- 
mation of the Western Society of Malacologists, 
but he evidently did not have a formal role with 
that organization (M. Woolsey to E. V. Coan, in 
litt., Dec. 26, 2007). 

Taylor also presented a paper at the 1964 
meeting of the American Society of Zoologists, 
and he (or his co-authors) presented papers 
at the 1957 annual meeting of the Geological 
Society of America, and the 1992 regional 
meeting of the Cordilleran Section of the Geo- 
logical Society of America. He also submitted 
short papers on the Physidae for malacologi- 
cal conferences in Zhytomyr, Ukraine and 
Vladivostok, Russia (Taylor, 2002a, 2004b), 
and attended the first meeting (Korniushin & 


186 KABAT & JOHNSON 


FIG. 7. At the First European Malacological Congress, London, 1962. Left to right: John B. Burch, 
Anne Gismann, Melbourne R. Carriker and Dwight W. Taylor (courtesy of John B. Burch). 


Melnychenko, 2002; V. Anistratenko to A. R. 
Kabat, in litt., May 30, 2007), but did not at- 
tend the second meeting (K. A. Lutaenko toA. 
К. Kabat, in litt., May 21, 2007). In 2004, Tay- 
lor made a second visit to Ukraine to collect 
freshwater snails in the region around Kiev. 

In the late 1970s, James McLean of the Los 
Angeles County Museum of Natural History 
saw Taylor at a meeting of the Geological So- 
ciety of America in San Diego. McLean, who 
had known him since the early 1960s, sug- 
gested that he should resume attending the 
meetings of malacological societies, since so 
few people in the field knew him. The response 
was to the effect that he “would rather not be 
known,” and that he did not like any of the 
people conducting research in freshwater 
malacology. 


Biogeographic History of the Snake River 


Some of Taylor’s major contributions to mol- 
luscan biogeography arose from his studies 
of the fossil and living mollusks of the Snake 
River in southern Idaho. The modern Snake 
River originates in northwestern Wyoming and 
flows westwards across southern Idaho be- 
fore turning sharply northwards through the 
Hells Canyon to Lewiston, where it turns west- 
wards again through southeastern Washing- 
ton and flows into the Columbia River. The 
Snake River currently drains most of south- 
ern Idaho, except for the Bear Lake region in 
the southeastern corner. 

In the late 1950s, Taylor made a reconnais- 
sance field trip to the Snake River Plain with 
Survey geomorphologist Harold Malde and 


DWIGHT WILLARD TAYLOR 187 


volcanologist Howard Powers. Later, when 
Malde was mapping the Glenns Ferry area, 
Taylor worked as his field assistant. These and 
other collecting trips to the Snake River re- 
gion resulted in numerous internal Survey re- 
ports, in which Taylor documented over 100 
molluscan species, and his reports were a pri- 
mary source of data for the key paper by Malde 
& Powers (1962) on the geologic history of the 
Snake River region. 

However, the fact that the Hells Canyon sec- 
tion represents a geologically much younger 
canyon than the eastern part of the Snake 
River led other researchers in the mid-1950s 
to hypothesize that the Snake River did not 
Originally drain north through the Hells Can- 
yon but instead flowed south through Nevada 
to the Sacramento region of central Califor- 
nia. These researchers correctly inferred that 
the Hells Canyon did not open until fairly re- 
cently. Taylor (1960a) instead hypothesized, 
based on the distribution of several species of 
freshwater mollusks, that during the Pliocene 
or early Pleistocene, the Snake River flowed 
southwest to southern Oregon, into the Kla- 
math Basin. He also hypothesized that the 
Bonneville Basin of northern Utah (today an 
isolated, landlocked basin) then drained into 
the Snake River Basin through the Bear Lake 
region (Taylor, 1960a). Later he determined 
that the species studied in his 1960 paper were 
exceptional, and that the more usual distribu- 
tion track instead ran from southern Idaho 
through southeastern Oregon into eastern 
California, but bypassing the Klamath Basin 
(Taylor, 1966b: 22, 27). This distributional 
track, which Taylor was to call the “fishhook” 
distribution (after its resemblance to a fishhook 
when drawn on а map), proved influential, with 
other freshwater organisms seemingly shar- 
ing a congruent distributional pattern. His sub- 
sequent research provided what appeared to 
be further confirmation for his novel hypoth- 
eses of the drainage history of the Snake River 
Basin (Taylor, 1985c; Taylor & Smith, 1981). 

Recently, Hershler & Liu (2004b: 933-935), 
based on genetic analyses of species of 
Pyrgulopsis (Hydrobiidae) from this region, 
instead concluded that the Snake River Basin 
and Oregon Lakes faunas were not separate 
monophyletic groups, as would be suggested 
by Taylor’s hypothesis in which the two fau- 
nas were originally connected before the 
Snake River changed its direction. Instead, 
they determined that the shared presence of 
species in both regions, and the weak genetic 


differentiation between the two regions, were 
probably a result of occasional Pleistocene 
overflow from one basin into the other, with- 
out having the Snake River itself flow into Or- 
egon. Thus, although Taylor’s hypotheses 
concerning the flow of the Snake River may 
no longer be supported, he did correctly rec- 
ognize a partially shared fauna between the 
Snake River Basin and the southern Oregon 
Lakes region, and his ideas stimulated exten- 
sive research into their complex drainage his- 
tory and biogeography. 

Taylor also studied the biogeography of other 
Pliocene-Pleistocene faunas, which resulted 
in a major paper on the “Blancan” nonmarine 
mollusks of the Late Pleistocene and early 
Pleistocene (Taylor, 1966b). This paper, which 
summarized all the published data, reviewed 
numerous museum records, and included ex- 
tensive comparisons with the fossil mammal 
and fish faunas of that period, also proved in- 
fluential in understanding the distribution and 
evolution of these faunas. G. R. Smith (to A. 
К. Kabat, in litt., June 19, 2007) identified sev- 
eral key biogeographical hypotheses from this 
paper, and his earlier paper (Taylor, 1960a), 
as follows: (1) showing a pattern of distribu- 
tion of past hydrographic connections does not 
imply that those connections all existed at the 
same time (Taylor, 1960a: 325, 332); (2) geo- 
logical changes have occurred more rapidly 
than species have evolved, so that early Pleis- 
tocene species have persisted in havens 
where geological changes did not take place 
while becoming extinct elsewhere (Taylor, 
1960a: 333); (3) “tectonic activity can affect 
the differentiation of mollusks in 2 ways: ei- 
ther directly, by separating areas of formerly 
continuous habitat or joining formerly isolated 
habitats; or indirectly by environmental 
changes,” the former resulting in speciation, 
and the latter by stimulating natural selection 
in changing environments (Taylor, 1966b: 14); 
(4) that the “Nebraskan” glaciation, formerly 
thought to initiate the Pleistocene, was not the 
first glacial event in the Great Plains region 
(Taylor, 1966b: 8-9); (5) the earlier glaciation 
events were less severe and less widespread 
than the later (Wisconsin) events (Taylor, 
1966b: 9-10); and (6) “species living in a vari- 
ety of habitats, or in a widely available habitat 
such as shallow or seasonal ponds” have 
broader temporal and geographical distribu- 
tions than those in geologically active and iso- 
lated areas (Taylor, 1966b: 16). Malde (1972: 
14-15) credited Taylor’s Blancan paper for its 


188 KABAT & JOHNSON 


“valuable discussions” of the molluscan spe- 
cies from the Glenns Ferry Formation (Plio- 
cene) of the Snake River, Idaho, but concluded 
that subsequent research cast doubt on his 
interpretations of the stratigraphic relations of 
this fauna. 

His extensive research on Snake River bio- 
geography led to his developing strong pro- 
fessional relationships with several non- 
malacological colleagues whose work he was 
to greatly respect. Although not as strong as 
his friendship with Hibbard, whom Taylor re- 
garded as his mentor and surrogate parent, 
and whose style and prejudices he adopted, 
these colleagues remained major influences. 
Robert Bright (1928-1995), a paleontologist at 
the University of Minnesota, and a native of 
southern Idaho, went on many field trips with 
Taylor in Idaho and Utah. Aldrich Bowler (1915- 
2007), a playwright, potter, and environmental 
activist who lived near Bliss, on the Snake River 
in southern Idaho (Ronayne, 2007), stimulated 
Taylor’s appreciation of environmental and 
conservation issues. Harold Malde (1923- 
2007), the Survey geologist, looked after 
Taylor's well-being from a distance, and helped 
arrange grant support from the McKenna Foun- 
dation. Taylor made two visits of several weeks’ 
duration to the Venezuela residence of Leon 
Croizat (1894—1982), the unconventional bio- 
geographer, whose theories on the distribution 
of animals directly influenced Taylor’s recon- 
struction of the biogeographic distribution of 
freshwater mollusks. (G. R. Smith to A. R. 
Kabat, in litt., June 19, 2007). 

Gerald (Jerry) Smith, a professor at the Uni- 
versity of Michigan who published extensively 
on fossil freshwater fishes of the Great Basin, 
and who made collecting trips with Taylor, was 
also an influence on Taylor, who later harshly 
criticized Smith’s biogeographical conclusions. 
Smith well remembers long conversations with 
Taylor as they drove across the American west 
in Taylor’s blue Volvo with a license plate that 
read “SNAIL.” Smith recognized that study of 
the Snake River and Great Basin faunas had 
to be guided by “Taylor’s methods and his prin- 
ciple that hydrologic, topographic, and climatic 
features change more rapidly than do lineages 
of organisms.” (Smith et al., 2002: 177). For 
that reason, Smith emphasized the importance 
of using biological “data that are independent 
of geological assumptions,” in order to avoid 
“the logical circularity inherent in using geo- 
logical events to date evolution and in using 
evolutionary differentiation to date geological 
events.” (Smith et al., 2002: 179). 


Hydrobiidae of Cuatro Ciénegas 


In 1961, Carl Hubbs (1894-1979), an ich- 
thyologist at the Scripps Institution of Ocean- 
ography, and an expert on the desert fishes of 
the Great Basin, sent Taylor some freshwater 
gastropods from the desert springs near 
Cuatro Ciénegas, in the Coahuila state of 
Mexico. This small desert basin contains nu- 
merous interconnected springs, lakes, and 
streams, with a complex drainage pattern, and 
its geographic isolation has resulted in numer- 
ous endemic species of fishes, turtles, and 
invertebrates, aptly described as a “new world 
for biologists” (Taylor & Minckley, 1966). Tay- 
lor recognized that “a novel fauna had been 
found” in the specimens sent by Hubbs, but 
that fieldwork was required to study these 
specimens. Hence, when Wendell Minckley 
(1935-2001), an ichthyologist at Arizona State 
University, later sent Taylor preserved speci- 
mens of “remarkable Hydrobiidae ... he offered 
an instantly accepted opportunity to visit 
Cuatro Ciénegas” (Taylor, 1966c: 155). Taylor 
then made several visits, and these collections 
resulted in his best-known paper, which de- 
scribed numerous new endemic species, gen- 
era, and subfamilies of Hydrobiidae (Taylor, 
1966c). This paper was to be the apex of his 
study of the Hydrobiidae. 

Taylor (1966c) described five new genera 
and twelve new species of Hydrobiidae from 
Cuatro Ciénegas, all endemic to that small 
desert basin, then the highest known diver- 
sity of hydrobiids from such a small region. 
Some of the new species were remarkable in 
being brightly colored, as most freshwater 
mollusks are drab. One, Paludiscala caramba, 
had strong shell sculpture like a marine snail, 
and was aptly named after Taylor's response 
to seeing the shell: he explained that “caramba” 
was “an exclamation, loosely translated from 
my original remarks at seeing the shells. An 
epitoniid-like snail in the arid interior of north- 
ern Mexico is thoroughly implausible” (Taylor, 
1966c: 208). 

José Parodiz (1911-2007), of the Carnegie 
Museum, was independently studying the 
same fauna, but ceased his research upon 
seeing Taylor's first paper. Parodiz, who was 
knowledgeable about the hydrobiids of South 
America (Kabat & Hershler, 1993: 76), pro- 
vided suggestions as to genera that could be 
compared with the Cuatro Cienegas fauna, but 
later noted that his comments were completely 
ignored. Instead, Taylor wrote to Parodiz, ad- 
mitting that “perhaps you will think | am too 


DWIGHT WILLARD TAYLOR 189 


much of a splitter — this question of ranking is 
always subject to personal interpretation,” and 
that “you can judge for yourself whether I’m 
crazy or not!” (D. W. Taylor to J. J. Parodiz, in 
litt., Oct. 10, 1966). Adecade later, Taylor rec- 
ognized the limitations of his research, noting 
that it “was necessarily hasty and somewhat 
superficial, lacking the morphological detail 
one would like.” (D. W. Taylor to R. Hershler, 
in litt., Sept. 18, 1978). He placed part of the 
blame on the “influence of J. P. E. Morrison, 
the Smithsonian curator who was (supposedly) 
a source of guidance when | went to Wash- 
ington, D.C., to work in 1955.” (Ibid). 

In order to determine the systematic rela- 
tionships of these new species and genera, 
Taylor had to diagnose and revise the North 
American subfamilies of Hydrobiidae, which 
led to the recognition of four new subfamilies 
(three endemic) and two new tribes (Taylor, 
1966c: 215). Thus, this paper was important 
not only for the descriptions of the endemic 
new taxa, but also for its comprehensive, and 
long-overdue, revision of the higher classifi- 
cation of the Hydrobiidae. Taylor continued to 
make collecting trips to this region, and his 
“serendipitous observation” of amphipods 
trapped in the surface film layer of the springs 
led to the discovery of new methods for col- 
lecting these minute crustaceans, and the 
naming of “Taylor's Spring” (Cole, 1984). In- 
explicably, he never published again on this 
fauna, despite his subsequent collections from 
that region. 


Departure from the Survey — the Wandering 
Years 


In early 1965, Taylor obtained a change of 
station posting that allowed him to work at the 
University of Michigan Museum of Zoology, 
while remaining on the Survey’s payroll. Burch 
arranged with the museum’s director, T. H. 
Hubbell, for Taylor to serve as an unpaid Re- 
search Associate. Since there was no free 
space in the Mollusk Division, Burch planned 
for Taylor to share an office with a graduate 
student, Charlotte M. Patterson, who was 
studying the Succineidae (Gastropoda: Pul- 
monata). However, Taylor refused to share the 
office, so that when Ms. Patterson came to 
the office the next day, she found all of her 
belongings on the floor in the hallway. Burch 
attempted to mollify her by providing her with 
table space in the Mollusk Division library. 

In May 1965, Taylor participated in a National 
Science Foundation-funded research trip to 


Japan to study freshwater gastropods, which 
actually resulted in a paper on freshwater 
bivalves co-authored with Teruya Uyeno, for- 
merly a graduate student at Michigan who had 
studied fossil fishes. (Taylor & Uyeno, 1966) 
(Fig. 8). This trip started out as a grant pro- 
posal written by Burch, which originally was 
to include two graduate students, George 
Davis and Charlotte Patterson. Shortly after 
the grant was approved, Davis received his 
Ph.D., and obtained his own funding from the 
U.S. Army to pursue similar research, thereby 
creating a vacancy in Burch's project. Hence, 
Burch asked Taylor if he was interested in 
participating, and he eagerly accepted. Since 
Burch, Davis, and Patterson each had their 
own research projects, they anticipated that 
Taylor would do his own research in Japan. 
To their surprise, Taylor immediately prepared 
a detailed research plan of operation, with 
numerous preparatory tasks that Burch, Davis, 
and Patterson were to conduct, all relating to 
Taylor’s own research. Burch had to explain 
to Taylor that they were already committed to 
do their own research, and would not be able 
to assist him with his own research; he did not 
appreciate their disinterest. As it happened, 
Taylor went to Japan separately from Burch, 
so their paths did not cross, and Burch never 
saw Taylor again. Burch later commented that 
Taylor could have made numerous contribu- 
tions to the malacology and paleontology re- 
search programs at Michigan, and could have 
learned much from his colleagues at Michi- 
gan, had he been more willing to work with 
others in a cooperative manner (J. B. Burch 
to А. К. Kabat, in litt., Aug. 10, 2007). Davis, 
who also recognized Taylor’s talents, remarked 
that he was too narrow in his approach to evo- 
lutionary issues, ignoring new developments 
in phylogenetic methodologies, biogeographic 
theories, and statistical approaches, thus re- 
stricting his conclusions as he was too highly 
focused on his own interests (G. M. Davis to 
A. R. Kabat, in litt., Nov. 14, 2007). 

In September 1965, Taylor transferred from 
Michigan to the Survey’s smaller office in 
Menlo Park, California, just north of Stanford 
and a short drive south of San Francisco. This 
transfer was met with relief by Taylor’s col- 
leagues at the Survey in Washington, D.C. As 
was Said to us, Dwight was known for burning 
his bridges while standing on them. 

However, the transfer did not improve his 
professional relationships. He wrote that, “Ac- 
cording to Tom Dutro there will be space for 
me there, but | have strong opposition from 


190 KABAT & JOHNSON 


FIG. 8. Taylor, during 1965 visit to Hokkaido, Japan. Photograph taken by Teruya Uyeno (courtesy of 


Gerald Smith). 


several people there and am laying alterna- 
tive plans — i.e., I’m going to be with the Sur- 
vey in California, or unattached and working 
independently on grants in California. We shall 
see.” (D. W. Taylor to W. К. Emerson, in litt., 
Mar. 8, 1965). 

In 1967, he resigned from the Survey and 
became an Associate Professor of Zoology at 
Arizona State University (ASU), where 
Minckley was on the faculty. The Zoology De- 
partment was founded in 1962, and Minckley 
took the lead in developing major research 
programs in the systematics, biogeography, 
and ecology of the local fauna (Collins et al., 
2002: 258). While at ASU, Taylor taught biol- 
ogy and zoology courses, including the inver- 
tebrate zoology course, which had four field 
trips to the Gulf of California. Taylor also su- 
pervised the Master’s thesis of Wesley Farmer, 
who was one year younger than Taylor. Farmer 
published his thesis in 1970, a review of swim- 
ming in marine gastropods. 

Taylor then agreed to supervise a second 
graduate student, Jerry Landye, who was in- 
terested in the freshwater mollusks of Cuatro 
Ciénegas, particularly Mexipyrgus. Landye, 
while a student at Washington State Univer- 


sity, had spent several field sessions with Tay- 
lor, during which he carefully mentored 
Landye, by showing him how to record eco- 
logical and geological data in the field, and how 
to collect and preserve specimens. Landye did 
extensive collecting at Cuatro Ciénegas, both 
on his own and with Taylor. Thus, Landye was 
looking forward to pursuing doctoral studies, 
commencing in the fall of 1969. However, af- 
ter only two years at Arizona State, Taylor evi- 
dently became frustrated with the academic 
bureaucracy, for he resigned his faculty posi- 
tion in the summer of 1969, shortly before 
Landye's arrival. Hence, Landye turned to the 
study of desert fishes and doing consulting 
work, before returning to the Hydrobiidae, 
years later. Taylor ended any further contacts 
with Landye by the mid-1970s. 

In the fall of 1969, Taylor moved to San Di- 
ego where he became an unpaid Research 
Associate at the San Diego Museum of Natu- 
ral History, operated by the San Diego Soci- 
ety of Natural History. While at San Diego, he 
published two annotated bibliographies on the 
Cenozoic freshwater mollusks of western 
North America, which remain essential refer- 
ences (Taylor, 1970, 1975). He also intended 


DWIGHT WILLARD TAYLOR 19 


to author a book on the terrestrial gastropods 
of California, which was not completed (Mar- 
tin, 1972: 9). He co-authored, with Allyn G. 
Smith, a biography of Harold Hannibal (1889- 
1965) that is interesting in its candid descrip- 
tion of Hannibal’s life: 

“Harold Hannibal spent a meteoric career in 
Tertiary paleontology, and in studying living 
and fossil freshwater mollusks, during the early 
years of this century. His brilliant early work 
was tragically cut short, and he passed more 
than half his life in a mental institution only a 
few miles from his birthplace. Today he is re- 
membered mainly through colorful stories dat- 
ing from the period of his illness.” (Taylor & 
Smith, 1971: 303). 

Other than the “mental institution,” those 
words could have been written about Taylor 
himself. 

It was at this time that Taylor generously 
donated “a substantial part of the publication 
costs” of the second edition of Myra Keen’s 
Sea Shells of Tropical West America, as Keen 
(1971: v) graciously acknowledged. 

Richard Squires, a professor of geology and 
paleontology at California State University — 
Northridge, tried to have Taylor serve as the 
outside reviewer for the master’s thesis of Ri- 
chard Vincent Lamb, who studied the fossil 
freshwater mollusks found in the La Brea Tar 
Pits in Los Angeles (Lamb, 1989). Alas, Lamb 
submitted his thesis proposal one day late, and 
Taylor rejected the proposal solely because it 
was late?. 

The San Diego position also proved short 
lived, for by 1974, he was at the Pacific Ma- 
rine Station, operated by the University of the 
Pacific, and located at Dillon Beach, north of 
San Francisco, evidently in an unpaid capac- 
ity. That affiliation lasted until the University of 
the Pacific announced that the marine station 
would be closed in 1979. Hence, in 1978, he 
moved a few miles to the Tiburon Center for 
Environmental Studies, operated by San Fran- 
cisco State University. The Tiburon Center also 
had two ichthyologists who were interested in 
biogeography: Tyson Roberts, formerly of 
Harvard's Museum of Comparative Zoology 
(MCZ), and Leonard Compagno, now at the 
South African Museum. That position also did 
not last long, since several publications in the 
early 1980s gave his mailing address as his 


home address in Tiburon. By 1985 he moved 
to the Department of Geology, Oregon State 
University, Corvallis, where he was to spend 
the rest of his career. The paleontological 
couple Arthur Boucot and Jane Gray, both on 
the faculty, recruited Taylor to come to Oregon 
as a research associate. 

While living in California, Taylor resumed at- 
tending gentlemen’s clubs. At one club, Martha 
Ruby Chavez, pretended to fall in love with 
Taylor, and “accepted” his marriage proposal. 
Taylor, who was 47, was so smitten with his 
prospective bride that she arranged for her 
“brother-in-law” to meet him a few days in ad- 
vance of the wedding. This señor convinced 
him that he should invest $50,000 in an orange 
grove in Mexico. He immediately wrote a check 
for that amount. The señor, the orange grove, 
and the money were never seen again (Taylor 
told others that it was “her father’s cattle 
ranch”). Nonetheless, he married her in Marin 
County on July 11, 1979. It appears that he 
never told his former colleagues that he had 
married, and they never met his wife. 

The marriage proved financially disastrous, 
and Taylor filed a petition to divorce his wife 
barely six months later, in January 1980, with 
the divorce finalized on June 9, 19826. Under- 
standably embittered, he later admitted to sev- 
eral colleagues that the marriage was a 
disaster, but never explained the details. 

In order to resolve his financial commitments 
made before or during the marriage, and to 
obtain funds to live on, he had to sell most of 
his sizable malacological library, retaining only 
key references on freshwater mollusks. James 
H. McLean of the Los Angeles County Museum 
of Natural History tried to interest the then- 
Director, Giles W. Mead, into purchasing the 
library on behalf of the museum, but even the 
wealthy Mead balked at the $200,000 asking 
price. Instead, in 1984, Taylor sold much of 
his library, comprising over 90 large boxes, and 
including complete runs of a number of mala- 
cological serials, through Richard Petit on a 
consignment basis. Alas, many of the rare an- 
tiquarian titles, including folio works such as 
Perry, Gualtieri, and d’Argenville, were previ- 
ously damaged or destroyed when his Tiburon 
office was flooded with muddy water from a 
landslide; Taylor was too distracted by the di- 
vorce to dry out those books. 


In 1996, Lamb earned his doctorate at the University of Michigan, under Burch, on reproduction in the snail Zonitoides 


nitidus. 


°In re the Marriage of Dwight W. Taylor, Petitioner, and Martha К. Chavez-Taylor, Respondent, No. 96232 (Superior Court, 


State of California, County of Marin). 


192 KABAT & JOHNSON 


Reports on Endangered Species 


From 1978 through 1987, Taylor authored a 
series of internal reports to federal government 
agencies, principally the U.S. Fish & Wildlife 
Service (USFWS), on the distribution and 
abundance of various freshwater mollusks that 
were candidates for listing as endangered or 
threatened species under the Endangered 
Species Act of 1974. Although Congress had 
passed this legislation, signed by President 
Nixon, the authority to designate a species as 
endangered or threatened, which could lead 
to restrictions on altering the habitat, was del- 
egated to the U.S. Department of the Interior, 
which, in turn, hired numerous field biologists 
on а contract basis to prepare these endan- 
gered species reports. Taylor’s detailed reports 
were influential to the sponsoring federal agen- 
cies and to other researchers, and some have 
been cited in the formal literature. These re- 
ports, and several others submitted to state 
agencies, are cataloged in a separate anno- 
tated bibliography (Kabat, in prep.). 

His November 1986 report to the USFWS, 
on the “Fish Springs pond snail,” Lymnaea 
(Hinkleyia) pilsbryi Hemphill, 1890, from Juab 
County, Utah, regretfully concluded that this 
species was now extinct, with all Known speci- 
mens being long dead. Taylor placed blame 
on those responsible for the extinction of this 
species: none other than his sponsor. He forth- 
rightly criticized the USFWS for management 
practices that favored waterfowl (ducks and 
other game birds) over snails, since enhanc- 
ing habitats for birds led to the extinction of 
this snail. Indeed, his report remarked, in all 
capital letters, that “BIRD FARMS ARE NOT 
NATURAL HABITAT.” It is not surprising that 
this was his last report for the USFWS’. 


Biogeographical Research in Oregon 


At Oregon State University, Taylor resumed 
his biogeographical studies, resulting in the 
publication of several wide-ranging syntheses 
of the historical distribution of freshwater mol- 
lusks. These articles reflected both his decades 
of research in systematics and biogeography, 
and his somewhat iconoclastic approach to 


these issues. Three papers from 1985, 1987, 
and 1988 merit further discussion. 

First, Taylor, at the invitation of Charles 
Smiley of the University of Idaho, participated 
in a symposium, Late Cenozoic History of the 
Pacific Northwest (June 3-7, 1979), which in- 
cluded a field trip to the Clarkia fossil beds of 
northern Idaho. The symposium volume con- 
tains a lengthy paper, “Evolution of freshwa- 
ter drainages and molluscs in western North 
America” (Taylor, 1985c). This comprehensive 
review of numerous drainage basins from 
Idaho to Texas concluded that the subdivision 
of drainages caused (or facilitated) speciation 
with most speciation occurring in the earlier 
part of the Tertiary but not in the Pleistocene, 
even though the Great Basin region had ex- 
tensive Pleistocene lakes. Yet, Taylor rejected 
any attempt to use phylogenetic analyses of 
these mollusks to reconstruct the geologic his- 
tory of this region. By way of explanation, be- 
ginning in the 1970s, some researchers 
combined cladistic (Hennigian) methods of 
phylogeny reconstruction with biogeographic 
analyses in an attempt to show that geologic 
events correlated with, or caused, speciation. 
Taylor would have none of this: 

“Phylogenetic interpretations thus play no 
part in these inferred drainage histories. There 
is no element of cladistic analysis, and none 
whatever of that abhorrence, ‘vicariance bio- 
geography.’ | have previously (Taylor, 1960) 
acknowledged, and do so once more herein, 
my great debt to the ideas of Léon Croizat.” 
(Taylor, 1985c: 266). 

Taylor appears to have fundamentally mis- 
understood the concept of vicariance bioge- 
ography, which is actually based in part on 
Croizat's work. Croizat was a controversial 
botanist from Venezuela, who self-published 
several volumes in the early 1960s on “panbio- 
geography.” These poorly written, almost in- 
comprehensible books, did inspire others, 
notably Gareth Nelson and Donn Rosen of the 
American Museum of Natural History, who 
showed how phylogenetic hypotheses could 
be applied to geographic regions. Thus, while 
Taylor’s (1985) work sets forth an invaluable 
compilation and analysis of the molluscan 
fauna of these drainage basins, his interpre- 


‘Taylor also noted in this USFWS report that Richard H. Russell, who published on this species (Russell, 1971), was “the 
person with the most nearly first-hand acquaintance with the living species, [who] disappeared from the University of 
Arizona more than ten years ago while still a graduate student. This is a literal statement. His major professor, Walter 
B. Miller, and fellow graduate student Carl Christensen, have attested to this astonishing occurrence in person. No one 
in the field of malacology has heard from or of him since.” Similarly, Taylor became increasingly reclusive, with many of 
his former colleagues no longer hearing from him by the 1980s. 


DWIGHT WILLARD TAYLOR 193 


tations of the geologic history were limited by 
his misunderstanding of biogeographic meth- 
odologies. 

Further, Taylor did not adopt Croizat’s sub- 
sequent revisions to his theories, including 
Croizat’s adoption of vicariance biogeography 
as expounded by Nelson, Rosen, and others. 
Taylor was skeptical of vicariance biogeogra- 
phy because of its association with cladistic 
theories of phylogenetic reconstruction, as he 
rejected the reliance of cladistics on the use 
of unweighted, derived characters. He instead 
believed that his expertise allowed him to 
weight characters and “to synthesize diverse 
morphological, ecological and geological in- 
formation to provide an ultimate hypothesis 
that would stand the test of time” (G.R. Smith 
to A. К. Kabat, in litt., June 19, 2007). Not sur- 
prisingly, he also rejected the MacArthur and 
Wilson theory of island biogeography, since 
he believed that that theory ignored historical 
geology through its focus on immigration and 
extinction of species within isolated regions 
(/bid.). 

Taylor’s second biogeographic paper in the 
mid-1980s was co-authored with Robert 
Bright, the University of Minnesota paleontolo- 
gist who did much to reconstruct the stratigra- 
phy of southern Idaho and northern Utah. This 
paper, “Drainage history of the Bonneville 
Basin” (Taylor & Bright, 1987), concluded that 
the flow of water between southern Idaho and 
northern Utah had changed directions several 
times from the Miocene to the Quaternary. 
This, when combined with habitat fragmenta- 
tion, resulted in speciation. Taylor and Bright 
provided a remarkably detailed discussion of 
the changes in the species composition of 
these faunas, but rejected the attempts by ich- 
thyologists to conclude that changes in river 
flow alone, and not habitat changes, caused 
changes in species distribution: “This expla- 
nation [of species distributions] is needed to 
emphasize the serious misuse of the concept 
in recent ichthyological literature (Minckley and 
others, 1986). Subjecting a biogeographic 
abstraction to palinspastic reconstructions as 
if it were a pre-Cambrian anorthosite is pure 


‘geopoetry.’ The suggestions by those authors 
(p. 600) that the pattern might have been sev- 
ered as long ago as 10 million years, or per- 
haps persisted after 4.5 million years, indicates 
complete lack of understanding of the concept 
and of the principles involved.” (Taylor & Bright, 
Е 

“The most striking feature of his [Smith’s] work 
is that the Bonneville Basin drained into 
Hudson Bay during the latest Miocene and 
Pliocene, as inferred from some of the mod- 
ern fishes, and, strangely, gastropods. This 
unique and unprecedented conclusion has 
been criticized already (Taylor, 1985). The fun- 
damental error is that the drainage connec- 
tion responsible for faunal similarities (these 
are quite real) is simply assumed to be a 
through-flowing stream.” (Taylor & Bright, 
1987: 255). 

It should be noted that the recipients of 
Taylor's criticisms were two ichthyologists with 
whom Taylor had formerly collaborated but had 
since fallen out with: Wendell Minckley of Ari- 
zona State University, who had worked with 
Taylor on the remarkable fauna of Cuatro 
Cienegas, Mexico (Taylor & Minckley, 1966); 
and Gerald R. Smith of the University of Michi- 
gan, who co-authored a paper with Taylor on 
Pliocene mollusks and fishes (Taylor & Smith, 
1981). Smith later explained this criticism: Tay- 
lor originally suggested this biogeographical 
connection to Smith, who incorporated it into 
a draft manuscript. However, when Taylor re- 
viewed the manuscript, he harshly criticized 
his own suggestion (!), upon which Smith wa- 
tered it down in the final, published version. 
Yet, it was the manuscript formulation, not the 
published version, which Taylor was to remem- 
ber and criticize (С. К. Smith to A. К. Kabat, in 
litt., June 20, 2007). 

Finally, in 1988, at the invitation of Jane Gray, 
Taylor wrote a lengthy article, “Aspects of 
freshwater mollusc ecological biogeography” 
(Taylor, 1988b). The primary value of this pa- 
per lies in its summary of the geographic dis- 
tributions of 42 families of freshwater and 
brackish-water mollusks. Taylor argued that 
species habitats and life history traits prima- 


ЗА “palinspastic reconstruction” is a paleo-geographic map showing the distribution of organisms or geological features on 
the continents as they were arranged in earlier geological eras. Anorthosite is an igneous rock common in pre-Cambrian 
formations, including the San Gabriel Mountains near the Webb School and Taylor's hometown. When the distribution of 
anorthosite is mapped onto a pre-Cambrian map of North America, the result approximates a straight line, indicating that 
these anorthosite outcrops were caused by a common geologic event. Taylor’s argument is that that while it may be 
suitable to map the distribution of rocks onto paleo-geographic maps, fossils were unsuitable for such mapping and 
analysis, as he believed that species expanded their ranges into suitable habitats, as opposed to geological events 


transporting species into new habitats. 


194 KABAT & JOHNSON 


rily determine the biogeographic distribution 
of each family, so that different families will 
have different distribution patterns. Thus, he 
once again rejected the attempts by other, 
unnamed malacologists (evidently including 
George Davis and Winston Ponder) to use 
geologic events to explain biogeographic dis- 
tributions: “The conclusions drawn herein dif- 
fer greatly from those in numbers of previous 
works. The differences are due to method. In- 
terpretations that fit the data of distribution to 
such transient views as the derivation of the 
South American fauna from North America, or 
the rafting of a fauna on the Indian plate, are 
fundamentally unscientific. Such interpreta- 
tions provide no test of anything except the 
will power of an individual author.” (Taylor, 
+9886: 513). 

Taylor did note, in his acknowledgments, that 
Ponder had reviewed the manuscript: “With- 
ering criticisms by Winston Ponder, of the Aus- 
tralian Museum, who disagrees in general and 
in particular with virtually everything in the 
paper, made me decide to discard it. Editor 
Jane Gray and Arthur Boucot extracted finally 
a revised paper from the reluctant author. The 
future will determine whether they merit cen- 
sure or praise; but anyway they have my 
thanks.” (Taylor, 1988b: 569). 

Thus, Taylor’s trilogy of papers from the 
1980s on freshwater molluscan biogeography 
remain useful for their detailed compilations 
of data on the geographical distribution and 
fossil records of these mollusks, notwithstand- 
ing the limitations in their methodology. The 
1985 paper is probably the most insightful, 
since it proposed a number of interesting bio- 
geographical hypotheses, some of which have 
been adopted by other authors. It should be 
possible for other researchers to analyze his 
data, suitably updated, using other method- 
ologies for reconstructing biogeographic dis- 
tributions and phylogenetic relationships. 

Taylor also participated in the study of a large 
collection of fossils obtained by Arthur Boucot 
and several other paleontologists from the 
Devonian of northwestern Saudi Arabia. Only 
one mollusk species was found, in high den- 
sities, in the zone between the Jauf and Jubah 
formations. He was able to determine that this 
was a nonmarine gastropod, referable to the 
Rissooidea “in the broad sense,” but it could 
not be identified to family or species (Taylor in 
Boucot et al., 1989: 558). This discovery is of 
significance in extending the fossil record of 
this superfamily from the Permian, and repre- 


sented “the earliest plausible freshwater 
snails,” thus suggesting that the sister-taxa of 
both the Rissooidea and the suborder 
Discopoda necessarily had to be equally an- 
cient (Taylor in Boucot et al., 1989: 558-559). 


New Mexico Paper on the Hydrobiidae (1987) 


While in Oregon, Taylor continued his re- 
search on the Hydrobiidae and other gastro- 
pods in artesian springs of New Mexico and 
Texas. When Taylor first published on the 
Hydrobiidae of northern Mexico in 1966, he 
essentially had that family to himself (as Jo- 
seph Morrison had mostly ceased publishing 
on hydrobiids), but within a few years, George 
Davis of the Academy of Natural Sciences of 
Philadelphia and Fred Thompson of the Florida 
Museum of Natural History began publishing 
on the anatomy and phylogeny of that family 
and others in the Rissooidea. 

By the late 1970s, Robert Hershler, a gradu- 
ate student of George Davis and Steven 
Stanley, began a major restudy of the anatomy 
and classification of the hydrobiids of Cuatro 
Ciénegas, as originally described by Taylor in 
1966 from the shells. Hershler’s thesis, which 
generously credited Taylor’s work while recog- 
nizing the limitations of his collecting methods 
and the few taxonomic characters used, re- 
sulted in significant changes, including synony- 
mizing some of Taylor’s new taxa (Hershler, 
1984, 1985). While Hershler was still in gradu- 
ate school, at Johns Hopkins University, he 
wrote to Taylor in an attempt to learn whether 
he was still interested in this fauna, and to dis- 
cuss his research. Taylor’s lengthy response 
told Hershler, in no uncertain terms, that since 
Taylor was still studying this fauna, it was “fun- 
damental and obligatory” that “there should be 
no overlap with [Taylor’s] present active study 
by anyone else” and that he would only dis- 
cuss possible thesis topics if Hershler chose to 
study snails or faunas that have “no one [else] 
studying them.” (D. W. Taylor to R. Hershler, in 
ИЕ, Sept. 18, 1978). When Hershler instead 
continued his research on the Cuatro Ciénegas 
fauna, Taylor then tried to interfere with 
Hershler’s graduate committee, upon which 
Stanley had to remind Taylor that any overlap 
in their research was scientifically appropriate, 
and requested, to no avail, that he should in- 
stead try to have “more comfortable interactions 
or even cooperation in the study of” this fauna. 
(S. M. Stanley to D. W. Taylor, in МЕ, Apr. 4, 
1979). 


DWIGHT WILLARD TAYLOR 195 


Taylor did not appreciate these new devel- 
opments, and viewed Hershler as yet another 
“young Turk” who was to be ignored in the 
hopes that he would go away. However, 
Hershler was hired at the Smithsonian, where 
he began publishing an extensive series of 
papers on the anatomy and phylogeny of the 
Hydrobiidae, some co-authored with Davis, 
Thompson, and others. 

Hershler, based on anatomical studies, de- 
termined that some of Taylor’s new species 
were either junior synonyms or assigned to 
the wrong genus, and that his subfamily clas- 
sification was not justified. Although Hershler 
(as had Davis and Thompson) tried to keep 
Taylor informed of their new publications, and 
Hershler even named a new genus and two 
new species after him, Taylor refused to com- 
municate with them. 

Over the decades, Taylor accumulated nu- 
merous specimens of Hydrobiidae and other 
freshwater gastropods from artesian springs 
and lakes in Texas, New Mexico, Arizona, 
Nevada, and California. He realized that 
Hershler was actively collecting these same 
faunas, so did not want to get scooped, and 
hastily arranged to have a manuscript with 
numerous new genera and species, mostly 
from New Mexico, described in a paper (Tay- 
lor, 1987) published in the Bulletin of the New 
Mexico Bureau of Mines & Mineral Resources, 
a journal not known for articles about fresh- 
water gastropods. Taylor deposited all the in- 
cipient types of his new species in the Los 
Angeles County Museum (LACM) and the 
University of Texas at El Paso, but refused to 
tell the curators at those institutions when his 
paper was published, and later refused to pro- 
vide them with any reprints, claiming that he 
could not afford reprints (although this New 
Mexico agency sells reprints for only $7.50). 
Notably, the bibliography of this paper did not 
cite any publications by Davis, Thompson, or 
Hershler, despite the direct relevance of their 
research. 

As a result, it was not until the end of 1987 
that Hershler and other malacologists found 
out about this research. Since Hershler already 
had a paper in press that described new spe- 
cies of Hydrobiidae from Arizona, he had to 
prepare a short addendum noting that three 
of the new species were junior synonyms of 
Taylor’s taxa; coincidentally, one was also a 
junior homonym (Hershler & Landye, 1988). 

In early 1988, Taylor wrote to Clif Coney 
(1949-1993), then the collections manager at 
LACM, to complain about Hershler’s work: 


“The person at the Smithsonian ... has taken 
or is in process of taking, the core of the mono- 
graph that | have been working on for the past 
12 years. The species represented by the 
types | have sent to the museum are or will all 
be preempted, except for those in the New 
Mexico publication and perhaps a couple of 
Physas now in press in Malacological Review. 
The bulk of those types you may discard as 
you wish.” 

“The study of snails was never anything | 
did for money, only for satisfaction and fun, 
and that motivation is all gone. Now my high- 
est priority, the overriding concern, is to make 
sure that under no possible circumstances will 
any of the results of my years of collecting, 
study, sorting and picking, measuring, preserv- 
ing, drawing, and paying for travel, supplies, 
typing, and illustration ever be of the slightest 
benefit or available in any way to that Phila- 
delphia type of malacologist.” 

“| won't be corresponding on this or any other 
subject in the future, and request that you de- 
lete my name from your mailing list and that 
of the museum.” (D. W. Taylor to C. Coney, in 
litt., Feb. 1, 1988). 

Sometime in the early 1980s, Taylor had 
deposited, in the LACM, 59 lots of incipient 
type specimens of new species to be de- 
scribed, nearly all in the Hydrobiidae. The an- 
notated list of these new species indicates that 
many were to be in new genera. Gale Sphon 
(1934-1995), then a curatorial assistant at 
LACM, assigned catalog numbers to these 59 
lots. Since only 21 of those species were de- 
scribed in the 1987 paper, and two were de- 
scribed elsewhere, that left 36 incipient types 
remaining as manuscript names. 

Although Taylor suggested in his 1988 letter 
that those latter specimens could be dis- 
carded, it is more likely that Coney arranged 
for their return to him, since they cannot now 
be found in the LACM. Unfortunately, there is 
no record in the LACM's files as to the dispo- 
sition of those specimens, and all the individu- 
als involved (Coney, Sphon, and Taylor) are 
now deceased (L. Groves toA. R. Kabat, pers. 
comm., Mar. 30, 2007). 

Since Hershler’s work on the Hydrobiidae 
continued unabated, Taylor never forgave him 
for having taken over as the expert on the 
North American fauna. In April 1992, when 
Taylor came to the MCZ for a week to study 
the extensive Physidae collection that Clench 
had accumulated, Hershler learned of this 
upcoming visit and called Boss to see if he 
could also come to the MCZ that week, so that 


196 KABAT & JOHNSON 


he could finally meet Taylor. Boss, remember- 
ing what had happened three decades previ- 
ously during Hubendick’s visit to the USNM, 
then called to obtain his approval. He was 
upset that Hershler might get to see him, and 
promptly told Boss that “If Hershler comes, I’m 
not coming!” Boss, desiring to have Taylor’s 
visit so that the MCZ’s sizable collection of 
Physidae could be recurated, reluctantly had 
to tell Hershler that he could not visit at the 
same time. Hershler never met Taylor at any 
other time. Even so, when Taylor came, he 
refused to engage in discussions with any of 
us present; coincidentally, Kabat was also vis- 
iting that same week from the Smithsonian’. 
He later complained to the MCZ administra- 
tion that the research grant provided to him 
was too small, another instance of the dog bit- 
ing the hand that feeds it. 


Monograph of Physidae (2003) 


His last two decades, after he abandoned 
the Hydrobiidae, were focused on research- 
ing the monograph of the Physidae, a family 
of freshwater pulmonates, which was pub- 
lished in March 2003. He previously described 
several new species of in this family, and be- 
gan a worldwide revision of this family, which 
had not been comprehensively analyzed in 
decades. Earlier authors, including Hubendick, 
tended to lump the species into only two or 
three genera. George Ang Te, the only other 
American who had studied the Physidae in the 
1970s while a graduate student at Michigan 
(Te, 1975, 1980), earned Taylor’s enmity for a 
misidentification, as he later wrote: “George 
Te, about whom | have a lot to say verbally 
but none of which should go on paper, was 
responsible for that misidentification.” (D. W. 
Taylor to К. D. Turner, in litt., June 25, 1983). 

In 1991, Taylor made his first of what be- 
came nearly annual visits to Costa Rica, where 
he did extensive collecting, made good use of 
the resources at the Museo de Zoologia de la 
Universidad de Costa Rica, San Jose, and the 
Instituto Nacional de Biodiversidad, Santo 
Domingo, Heredia (INBio), and helped the stu- 
dents with their malacological research 
(Barrientos & Springer, 2007: x-xi). Taylor’s 
enthusiasm for the local molluscan fauna knew 
no bounds: in addition to publishing a prelimi- 


nary checklist of the freshwater mollusks of 
Costa Rica (Taylor, 1994), he also intended to 
publish a guide to the freshwater molluscan 
genera of Costa Rica, a monograph on the 
mollusks of Colombia, and another monograph 
on the freshwater and brackish-water mollusks 
of Costa Rica, but these and other manuscripts 
remain unfinished (Barrientos & Springer, 2007: 
xi). Taylor evidently enjoyed his visits to Costa 
Rica, and his interactions with the local stu- 
dents, who fondly called him “Don Guillermo.” 
Alas, on one trip, a suitcase containing numer- 
ous specimens of Polymesoda (Corbiculidae), 
carefully collected from around the country, was 
stolen, presumably by thieves who thought that 
a heavy suitcase must contain objects of great 
monetary value. Taylor was able to laugh about 
this in later years, notwithstanding the difficulty 
of trying to re-collect this material (/bid.). Dur- 
ing these trips, Taylor documented his travels 
and commented on his colleagues by writing 
notes on postcards which he mailed to his 
home, as a diary; these postcards are an in- 
teresting record of his views (Ibid.). 

Taylor's research on the Physidae was long 
overdue, and his 2003 monograph (already out 
of print, although an online version is planned 
for 2008), included an invaluable catalogue of 
the over 400 names introduced in this family, 
and a summary of the known museum hold- 
ings of type specimens. His monograph was 
based on anatomical study of numerous spe- 
cies from around the world, including many 
previously only known from empty shells. He 
described 17 new species, bringing the total 
number of Recent species to about 80, and 
described 11 new genera and five new tribes 
to accommodate the morphological diversity 
in this family, which now encompassed 23 
genera. Some of the new taxa inadvertently 
first appeared as nude names in the proceed- 
ings of a malacological conference in Ukraine 
(Taylor, 2002a), but these names are not avail- 
able under Articles 13 and 16 of the Interna- 
tional Code of Zoological Nomenclature. Roth 
(2003), in a perceptive review, commended 
Taylor for examining numerous specimens and 
his detailed anatomical studies. 

However, two major limitations of this re- 
search, as Roth (2003) also noted, are that 
Taylor's anatomical characters were primarily 
or entirely limited to the “terminal male repro- 


“Taylor also visited the MCZ in the summer of 1990 for a brief examination of the Physidae. The two authors of this paper 
were the only ones present in the Mollusk Department during that visit, and he vociferously berated one of us (Kabat) 
for not showing up in the morning one hour earlier than the arranged time for his visit. 


DWIGHT WILLARD TAYLOR 197 


ductive system” (i.e., the penial complex), and 
that the classification methodology was sus- 
pect since of the “12 binary penial complex 
characters ... only six of the 12 provide any 
information for grouping; the others are 
autapomorphic [unique] for only one of the 23 
genera analyzed” (Roth, 2003: 365). Since the 
Physidae are hermaphroditic, Taylor could 
have obtained characters from the rest of the 
reproductive tract, or other organs and sys- 
tems. 

Moreover, his single, fully resolved phyloge- 
netic tree was not constructed using concepts 
of parsimony. Roth (2003: 365), using cladis- 
tic analysis of Taylor’s data, generated mul- 
tiple, equally parsimonious but not fully 
resolved trees, indicating that Taylor’s single 
fully resolved tree is misleading. 

A third limitation of Taylor’s research is that 
his attempts to integrate phylogeny with bio- 
geography are suspect, due to his continued 
misinterpretations of Croizat’s theories. Taylor 
concluded that because the most primitive spe- 
cies and genera are now found on the Pacific 
coast of Central America, this region had to be 
where the family originated and initially diver- 
sified (Taylor, 2003: 2, 18; 2004a: 279). As Roth 
(2003: 364) noted, this conclusion is question- 
able since it would require “a very long period 
of relative evolutionary stasis ... for those taxa 
Taylor regards as primitive,” with evolutionary 
novelties instead (or only) occurring away from 
the centers of origin. 

Despite these methodological limitations, his 
monograph of the Physidae is a commend- 
able effort to obtain data on the geographic 
distribution and male reproductive anatomy of 
this family. Subsequently, several researchers 
have begun reanalyzing the phylogeny and 
systematics of the Physidae based on molecu- 
lar techniques (Rogers & Wethington, 2007; 
Wethington & Lydeard, 2007). 


Taylor’s Mollusk Collections 


The disposition of Taylor’s extensive re- 
search collections of mollusks has not hith- 
erto been documented in the literature. 

First, in December 1984, he donated his col- 
lections of Unionidae (Bivalvia) and various 
brackish-water mollusks, particularly the genus 
Cerithidea (Potamididae), to the Los Angeles 
County Museum (LACM), partly because the 
then-collection manager, Clif Coney, was also 
interested in freshwater bivalves. As Taylor 
published little on those groups, that collection 
was probably of the least interest to him. 


Second, the University of Minnesota, Bell 
Museum of Natural History, acquired the 
“Dwight W. Taylor Collection of Mollusks,” com- 
prising 5,000 to 10,000 dry lots, and an unde- 
termined number of preserved specimens, 
primarily from the western U.S.A. Robert 
Bright, a professor at Minnesota and curator 
in the Bell Museum, was well known for his 
research on the fossil plants, pollen, and mol- 
lusks of southern Idaho and northern Utah 
(Davis, 1995). Bright made several field trips 
with Taylor, and they coauthored a paper (Tay- 
lor & Bright, 1987) on the Bonneville Basin. 
As a result, Bright was instrumental in having 
the Bell Museum acquire this collection. Yet, 
no effort was made to alert the malacological 
community to this important collection. It was 
not until after Taylor’s death that one of us 
(Kabat) came across the obituary of Bright that 
mentioned, in passing, Bright’s acquisition of 
this collection, identified as “one of the most 
significant in North America for western mate- 
rial” (Davis, 1995). 

Third, Taylor’s will arranged to have the re- 
mainder of his collection placed at the Museo 
de Zoologia, Universidad de Costa Rica (San 
Jose, Costa Rica). However, Yolanda Cama- 
cho-Garcia, the curator at that institution, ar- 
ranged with the California Academy of Sciences 
(CAS) to have the North American specimens 
deposited in the CAS, with the University of 
Costa Rica receiving the neotropical speci- 
mens. The originals of Taylor’s field notes were 
also divided between these two institutions, 
corresponding to the distribution of the speci- 
mens. However, the CAS also received a pho- 
tocopy of his field notes for the neotropical 
collections. The CAS notes are archived as 
“DW Taylor Colln, Acq #548, SLF T-13, Station 
Data.” Wendell P. Woodring, who also gave his 
extensive book and reprint library to the Uni- 
versity of Costa Rica in the early 1980s, may 
have inspired Taylor’s gift to that institution. 
Taylor retained type material of some of his new 
species in his personal collection; if not subse- 
quently deposited in museum collections, these 
specimens should now be with the material de- 
posited in the CAS in 2006. 


Last Years 


After completing the Physidae book in 2003, 
Taylor only published one further paper, a short 
synopsis of that book for a malacological se- 
rial in Uruguay (Taylor, 2004a), and submitted 
an abstract for a malacological meeting in 
Vladivostok that he was unable to attend (Tay- 


198 KABAT & JOHNSON 


lor, 2004b). His physical health deteriorated, 
forcing him to cease his research. Despite the 
financially disastrous divorce, Taylor must 
have achieved some financial stability in later 
years. Some measure of his finances is docu- 
mented by his contributions to candidates for 
U.S. President and Congress, as recorded on 
the website of the Federal Election Commis- 
sion, which identifies a total of $44,228 in do- 
nations from 2002 through 2006 (under federal 
law, individuals are limited in donations that 
can be made to any one candidate). With one 
exception, all of his donations were made to 
Democrats or Independents. What is surpris- 
ing in light of Taylor’s prior personal relation- 
ships, is that more than half of his donations 
($27,153) were made to 22 female candidates, 
such as senators Hillary Clinton (New York) 
and Barbara Boxer (California), with his 
smaller donations to six male candidates, in- 
cluding John Kerry and Howard Dean (unsuc- 
cessful Presidential candidates), and most of 
the rest to liberal political action committees"°. 


Afterword 


On August 3, 2006, Dwight Taylor died from 
cancer, at the Wayne Morse Ranch House, 
Eugene, Oregon. He is survived by two sis- 
ters, Sarah Taylor Stephenson of Greenwich, 
Connecticut, and Margaret (Maggie) Taylor 
Cunningham, an Episcopalian priest in Bryn 
Mawr, Pennsylvania. Dwight also had four 
nieces and two nephews, some of whom he 
never met. Several days before he died, he 
made a major donation to the Alf Museum. His 
will left the balance in his bank account to the 
Alf Museum, and arranged to have his Oregon 
house sold, with the proceeds going to the 
Webb School. 

On March 30, 2007, a memorial service was 
held at the Vivian Webb Chapel of the Webb 
School, followed by a lunch and informal re- 
marks in the Alf Museum, in front of the photo 
showing the Webb School's three finalists in 
the Westinghouse competition (Fig. 6). The 
religious service was conducted by Dwight's 
sister, Reverend Cunningham, who candidly 
admitted that he was not religious, so the ser- 


vice reflected his environmental concerns, in- 
cluding a recitation of the “Earth’s Ten Com- 
mandments” by Ernest Callenbeck and a 
homily from “A Sacred Place to Dwell: Living 
With Reverence Upon the Earth” by Henryk 
Skolimowski (Anonymous, 2007). 

After the lunch, Dwight’s sister Maggie led 
the remarks, noting with grief and profound 
regret that he had become estranged from his 
family. Malcolm McKenna then spoke, and the 
years rolled away as he recalled numerous joint 
collecting trips and their shared enthusiasm for 
the biogeography of mollusks and mammals. 
McKenna noted that they made good use of 
the collection and library of John Q. Burch. 
McKenna noted, with regret, that Dwight broke 
off contact with him and other paleontologists 
by the late 1960s. McKenna then called on one 
of us (Kabat) to discuss Dwight's malacologi- 
cal accomplishments and relationships. Don 
Lofgren, the Director of the Alf Museum, and 
Susan Nelson, the head of the Webb School, 
recounted Dwight’s visits to the Webb School 
and the Alf Museum, including his interests in 
conservation issues on the campus, and in the 
current activities at the Museum. 

Two of Dwight’s classmates, Guilford Bab- 
cock and Pete Akin, shared their friendly remi- 
niscences of Dwight. Professor Babcock 
discussed how he and his wife enjoyed sev- 
eral visits to Dwight in the past few years, and 
even though they did not agree on political is- 
sues, they had respectful discussions. Akin 
recalled a post-graduation trip that Dwight, two 
other classmates, and he took on old Highway 
99 throughout California, north to Crater Lake, 
and back south on the coastal Highway 1 to 
Newport Beach. Following these remarks, and 
a tour of the Alf Museum, Dwight's ashes were 
scattered in a ravine on the north end of cam- 
pus, where, in 1999, Alf's ashes were also.scat- 
tered. 

In conclusion, Dwight Taylor was a malacolo- 
gist and paleontologist who achieved numer- 
ous significant accomplishments in the 
taxonomy and biogeography of freshwater 
mollusks. Unfortunately, his inability to take 
criticism of his work, or to work with others, 
has impaired his professional reputation. 


Even stranger is his donation in 2003 to Lyndon H. LaRouche, a perennial fringe candidate for President. LaRouche was 
convicted т 1988 for conspiracy to commit mail fraud, and conspiracy to defraud the Internal Revenue Service (IRS), 
based on illegal fundraising during his 1980 and 1984 presidential campaigns, and served over a decade in federal prison. 
United States v. LaRouche, 896 F.2d 815 (4th Cir. 1990). LaRouche resumed his campaigns upon his release from prison, 
with even less success. Taylor’s donation to LaRouche is difficult to reconcile with his other donations, but may reflect 


his maverick approach to life. 


DWIGHT WILLARD TAYLOR 199 


ACKNOWLEDGMENTS 


We thank the following individuals who kindly 
shared their reminiscences of Dwight Taylor: 
Pete Akin, Guilford Babcock, Kenneth J. Boss, 
John B. Burch, Margaret Taylor Cunningham, 
George M. Davis, J. Thomas Dutro, Jr., Will- 
iam K. Emerson, Jerry Landye, Don Lofgren, 
Harold Malde, James H. McLean, Susan 
Nelson, Richard E. Petit, Barry Roth, and 
Gerald R. Smith. Kraig K. Adler provided infor- 
mation on Claude Hibbard as a mentor and 
teacher at Michigan. Robert Hershler provided 
invaluable insights into Taylor’s research, and 
allowed us to examine his copies of Taylor’s 
reports to government agencies, many of which 
John and Mary Lou Pojeta had obtained from 
the archives of the U.S. Geological Survey. 

John B. Burch, George M. Davis, Robert 
Hershler, James H. McLean, John Pojeta, Ri- 
chard E. Petit, Gerald R. Smith, and an anony- 
mous reviewer provided helpful comments on 
earlier drafts of this paper. Vitalij Anistratenko 
provided information on the malacological con- 
ference in Ukraine in 2002, and on Taylor’s 
second visit to Ukraine in 2004. Konstantin A. 
Lutaenko provided information about the ma- 
lacological conference in Vladivostok in 2004. 

Warren Blow (USNM), Lindsey Groves 
(LACM), Jann Thompson (USNM), and Paul 
Valentich-Scott (SBMNH) allowed Kabat to 
examine the Taylor type specimens in their 
collections, and provided further information 
on the curation of those specimens. Jean F. 
DeMouthe (CAS) and Yolanda Camacho- 
Garcia (UCR), through Eugene V. Coan, pro- 
vided information on Taylor’s type specimens 
in their collections. John Burch, Don Lofgren, 
Gerald R. Smith, and Paul Valentich-Scott pro- 
vided electronic versions of most of the pho- 
tographs of Taylor reproduced herein. Carla 
Dietrich expertly prepared these figures for 
publication. 


CATALOG OF TAYLOR’S NEW TAXA 
Acronyms and Abbreviations 
ANSP Academy of Natural Sciences of 


Philadelphia, Philadelphia, Pennsyl- 
vania 


BMNH The Natural History Museum, Lon- 
don, United Kingdom [formerly Brit- 
ish Museum (Natural History)] 

CAS California Academy of Sciences, San 


Francisco, California 


FLMNH Florida Museum of Natural History, 
Gainesville, Florida 

Instituto Nacional de Biodiversidad, 
Santo Domingo de Heredia, Costa 
Rica [mollusk collection now in the 
UCR, and curated as “MZUCR-INB”] 
Los Angeles County Museum of 
Natural History, Los Angeles, Cali- 
fornia 

Los Angeles County Museum of 
Natural History, Invertebrate Pale- 
ontology, Los Angeles, California 
Museum of Comparative Zoology, 
Harvard University, Cambridge, 
Massachusetts 

Museo de Zoologia, Universidad de 
Costra Rica, San José, Costa Rica 
[cited by Taylor as “UCR”] 
MZUCR- Museo de Zoologia, Universidad de 
INB Costa Rica, San José, Costa Rica, 
collections formerly housed in the 
Instituto Nacional de Biodiversidad, 
Santo Domingo de Heredia, Costa 
Rica [cited by Taylor as “INBio”] 

San Diego Society of Natural His- 
tory, San Diego, California 

UC University of Colorado Museum, 
Boulder, Colorado 

University of California Museum of 
Paleontology, Berkeley, California 
Museo de Zoologia, Universidad de 
Costa Rica, San José, Costa Rica 
[collections now curated at “MZUCR”] 
Museum of Zoology, University of 
Michigan, Ann Arbor, Michigan 
National Museum of Natural His- 
tory, Smithsonian Institution, Wash- 
ington, D.C. [formerly U.S. National 
Museum] 

University of Texas at El Paso, El 
Paso, Texas 

Departamento de Zoologia, Institu- 
to de Biologia, Universidad Nacio- 
nal Autonoma de México, Mexico 
City, Mexico 

Co. County 


INBio 


LACM 


LACMIP 


MCZ 


MZUCR 


SDSNH 


UCMP 


UCR 


UMMZ 


USNM 


UTEP 


ZIBM- 
CMNO 


Unless otherwise indicated, the type locality 
of new species is in the United States of 
America. Type specimens indicated as 
“USNM” are in the Recent Mollusk collection, 
except for fossil species, which are housed in 
the Invertebrate Paleontology collection. Al- 
though the types of Taylor’s 1987 New Mexico 
publication were deposited solely in LACM and 
UTEP, Artie Metcalf, curator of mollusks at 


200 KABAT & JOHNSON 


UTEP, arranged in 1990 to have some para- 
types shared with the USNM, ANSP, and 
FLMNH. Taylor’s types at Stanford University 
were transferred to the California Academy of 
Sciences in 1977 (Smith, 1978: 1). 

As a paleontologist, Taylor routinely provided 
detailed information on the type localities for 
new species, including the township and sec- 
tions for those species from the United States. 
He explained this system as follows: “Land in 
the United States is commonly divided into 
‘townships,’ each 6 miles square, that are num- 
bered according to tier (T.) and range (R.) from 
standard base lines and meridians. A town- 
ship is divided into 36 ‘sections,’ each section 
(sec.) 1 mile square.” (Gregg & Taylor, 1965: 
109). This information allows for precise de- 
termination of the type localities with reference 
to the U.S. Geological Survey topographical 
maps of the United States. 

Taylor described 132 new taxa (including two 
replacement names for junior homonyms), of 
which 54 (40.9%) were in the Hydrobiidae 
sensu lato, 39 (29.5%) in the Physidae, 36 
(27.3%) in other gastropod families (primarily 
Lymnaeidae, Planorbidae, Pleuroceridae, and 
Pupillidae), and 3 (2.3%) in the Bivalvia. These 
132 new taxa encompassed 12 family-level 
taxa (including 6 in the Hydrobiidae sensu lato 
and 5 in the Physidae), 31 genus-level taxa 
(including 12 in the Hydrobiidae sensu lato and 
11 in the Physidae), and 89 species (includ- 
ing 35 in the Hydrobiidae sensu lato and 24 in 
the Physidae). Some authors have recently 
proposed that the Hydrobiidae be divided into 
several families, e.g., Amnicolidae, Cochliop- 
idae (Wilke et al., 2001); however, since most 
of Taylor’s taxa have not been analyzed with 
respect to these families, we have used Hydro- 
biidae sensu lato for convenience. 

In 1988, Taylor described two new species 
of Physidae — Physa megalochlamys from 
Wyoming, and Р natricina from the Snake 
River, Idaho. The original description (Taylor 
1988) specified that the holotypes were de- 
posited in LACM, i.e., LACM 2255 and LACM 
2256, respectively, the numbers assigned by 
Gale Sphon when Taylor deposited the incipi- 
ent types. Since LACM 2256 had been inad- 
vertently used for the type specimen of another 
author’s new species, the holotype of P. 
natricina was recatalogued as LACM 2970 in 
2003. However, the type specimen of P. 
megalochlamys cannot now be found in the 
LACM, and it may have been returned to Tay- 
lor with the remaining, undescribed incipient 


types (L. Groves to À. К. Kabat, pers. comm., 
Mar. 30, 2007). Taylor (2003: 164) later wrote 
that the holotype of Р megalochlamys was 
CAS 114779, which may be the specimen re- 
moved from the LACM and subsequently de- 
posited in the CAS. 

Taylor described five new tribes, 11 new 
genera, and 17 new species of Physidae in 
his 2003 monograph. He submitted a short 
paper for the proceedings volume of a mala- 
cological meeting held on May 13-15, 2002, 
in Ukraine, which used many of these new 
names, without formal description (Taylor, 
2002a). ICZN Article 16.1 requires that “every 
new name published after 1999 ... must be 
explicitly indicated as intentionally new,” but 
Taylor’s 2002 paper does not specify which of 
the names are new, so they remained nude 
names until the publication of his 2003 mono- 
graph. Some of the names in the 2002 publi- 
cation also do not satisfy several other 
provisions of the ICZN, including Article 13.1 
(which requires a description for each new 
taxon); Article 13.3 (which requires a type spe- 
cies for each new genus); and Article 16.4 
(which requires type specimen(s) for each new 
species). 


adamantina, Tryonia (Paupertryonia) — Taylor, 
1987: 41—42, fig. 20. Type locality: Diamond 
Y Spring, Pecos Co., Texas. Holotype LACM 
2089; paratypes UTEP 10060; ANSP 
376031; FLMNH 160949; USNM 854075. 
Gastropoda: Prosobranchia: Hydrobiidae. 
Notes: Hershler (2001: 16-17) redescribed 
this taxon and transferred it to Pseudotryonia 
Hershler, 2001. 

alamosae, Tryonia (Paupertryonia) — Taylor, 
1987: 42-44, fig. 21. Type locality: Ojo 
Caliente, 700 ft W, 1,700 ft S, sec. 31, Т. 8$ 
R. 7W, Socorro Co., New Mexico. Holotype 
LACM 2188; paratypes UTEP 10061; USNM 
854701, 854702. Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Hershler 
(2001: 15-18) redescribed this taxon, which 
he made the type species of Pseudotryonia 
Hershler, 2001. 

albiconica, Limnaea (Stagnicola) — Taylor, 
1957b: 657-659, figs. 1-3. Type locality: 
White Cone Peak, sec. 12, T. 25N, R. 21E, 
1.5 mi S of White Cone Trading Post, Na- 
vajo Co., Arizona. White Cone local fauna, 
Hemphillian Age, Bidahochi Formation, 
Middle Paleocene. Holotype USNM 562081; 
paratypes USNM 562082. Gastropoda: 
Pulmonata: Lymnaeidae. 


DWIGHT WILLARD TAYLOR 201 


alfi, Helminthoglypta — Taylor, 1954c: 76-77, 
pl. 20, figs. 30-32. Type locality: Barstow Hills, 
7 mi N of Barstow, San Bernadino Co., Cali- 
fornia (“a series of three outcrops of a stra- 
tum of whitish volcanic ash lying in a straight 
line in the NW corner of Rainbow Basin”). 
Barstow Formation, Upper Miocene. Holo- 
type: CAS 70413 (ex Stanford Univ. Paleon- 
tological Type Collection 8075); paratypes 
CAS 70414 (ex Stanford Univ. 8076); SBMNH 
35323 (ex S. S. Berry collection 15373); 
SBMNH 119935 (ex S. S. Berry collection 
19935); LACMIP 4922, 4923 (ex W. O. Gregg 
collection 5925); D. W. Taylor collection 1107, 
1108, 1417, 1439, 3137, and 3138; and 
USNM 561450. Gastropoda: Pulmonata: 
Helminthoglyptidae. 

Amecanauta Taylor, 2003: 72. Type species 
Amecanauta jaliscoensis Taylor, 2003; origi- 
nal designation. Gastropoda: Pulmonata: 
Physidae. Notes: Taylor (2002a: 25) previ- 
ously used this name as a nomen nudum. 

Amecanautini Taylor, 2003: 72. Tribe in the 
subfamily Aplexinae, for Amecanauta Tay- 
lor, 2003, Mexinauta Taylor, 2003, Mayabina 
Taylor, 2003, Tropinauta Taylor, 2003, and a 
“name uncertain group” in the subfamily 
Aplexinae. Gastropoda: Pulmonata: Physi- 
dae. Notes: Taylor (2002a: 25) previously 
used this name as a nomen nudum. 

Apachecoccus Taylor, 1987: 32. Type species 
Apachecoccus arizonae Taylor, 1987; origi- 
nal designation. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: Hershler (1994: 5, 13) 
concluded that this taxon was a junior syn- 
onym of Pyrgulopsis Call & Pilsbry, 1886. 

“Aplexini” Taylor, 2003: 49. Tribe in the sub- 
family Aplexinae, for Amuraplexa Staro- 
bogatov etal., 1989, Paraplexa Starobogatov, 
1989; Aplexa Fleming, 1820, and Sibirenauta 
Starobogatov & Streletskaya, 1967. Gas- 
tropoda: Pulmonata: Physidae. Notes: Staro- 
bogatov (1967: 289) had already established 
Aplexinae as a subfamily, so Taylor is not to 
be credited as the author of this tribe, pursu- 
ant to ICZN Article 36.1 (“A name established 
for a taxon at any rank in the family group is 
deemed to have been simultaneously estab- 
lished for nominal taxa at all other ranks in 
the family group ... The name has the same 
authorship and date at every rank.”). 

Archiphysa Taylor, 2003: 177-178. Type spe- 
cies Physa lordi Baird, 1863; original desig- 
nation. Gastropoda: Pulmonata: Physidae. 
Notes: Taylor (2002a: 25) previously used 
this name as a nomen nudum (as “Archi- 
physe” [sic]). 


arizonae, Apachecoccus — Taylor, 1987: 32- 
34, fig. 15. Type locality: unnamed spring on 
N side of Gila River, about 2 mi N of Bylas, 
ЗА: 22E; 25,000. W and 15500ft:N 
of township line, Graham Co., Arizona. Ho- 
lotype LACM 2203; paratypes UTEP 10050; 
ANSP 376020; FLMNH 160939; USNM 
854090. Gastropoda: Prosobranchia: Hydro- 
biidae. Notes: Pyrgulopsis sancarlosensis 
Hershler in Hershler & Landye, 1988: 35- 
41, is a junior subjective synonym of Apache- 
coccus arizonae Taylor, 1987 (fide Hershler 
& Landye, 1988: 58). Hershler (1994: 18) 
recognized this taxon as a valid species of 
Pyrgulopsis Call & Pilsbry, 1886. 

ashmuni, Archiphysa — Taylor, 2003: 178-180, 
pl. 10, figs. 1-3, map fig. 176. Type locality: 
Ojo de Gallo, 1,650 ft N, 4,150 ft E, sec. 3, T. 
10N, К. 10W (35°07’20”М, 107°52’32”\М), 
Cibola Co., New Mexico. Holotype CAS 
146087; paratypes CAS 146088; BMNH 
2001307. Gastropoda: Pulmonata: Physidae. 

Austrinauta Taylor, 2003: 43—45. Type species 
Physa elata Gould, 1853; original designa- 
tion. Gastropoda: Pulmonata: Physidae. 
Notes: Taylor (2002a: 25) previously used 
this name as a nomen nudum. 

Austrinautini Taylor, 2003: 43. Tribe in the sub- 
family Aplexinae, for Austrinauta Taylor, 2003 
and Caribnauta Taylor, 2003. Gastropoda: 
Pulmonata: Physidae. Notes: Taylor (2002a: 
25) previously used this name as a nomen 
nudum. 

bernardinus, Yaquicoccus — Taylor, 1987: 34— 
35, fig. 16. Type locality: spring, 2,300 ft E, 
4,600 ft S of NW corner, section 15, T. 24S, 
R. 30E, Cochise Co., Arizona. Holotype 
LACM 2186; paratypes UTEP 10066; ANSP 
376019; FLMNH 160942, 160934; USNM 
854078, 854088. Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Pyrgulopsis 
cochisi Hershler in Hershler & Landye, 1988: 
41, is a junior subjective synonym of Yaqui- 
coccus bernardinus Taylor, 1987 (fide Hersh- 
ler & Landye 1988: 58). Hershler (1994: 21) 
recognized this taxon as a valid species of 
Pyrgulopsis Call & Pilsbry, 1886. 

blakeana, Pyrgulopsis — Taylor, 1950: 30-31, 
figs. 4-6. Type locality: shore of Salton Sea, 
by Fish Springs, Imperial Co., California. 
Quaternary. Holotype: SBMNH 35500 (ex S. 
S. Berry collection 13251); paratypes 
SBMNH 35501 (ex S. S. Berry collection 
13253); SBMNH 35502 (ex S. S. Berry col- 
lection 13259); SBMNH 35322 (ex S. S. 
Berry collection 13253); USNM 613966. 
Other paratypes “promised” to the collections 


202 KABAT & JOHNSON 


of W. O. Gregg, SDSNH, Stanford Univer- 
sity, and UMMZ. Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Taylor (1966c: 
196) subsequently concluded that this was 
a junior synonym of Tryonia protea (Gould, 
1865): 

brunei, Tryonia (Paupertryonia) — Taylor, 1987: 
44-45, fig. 22. Type locality: outflow of Phan- 
tom Lake Spring, Joe Kingston Ranch, Jeff 
Davis Co., Texas. Holotype LACM 2251; 
paratypes UTEP 10062; USNM 854631, 
854632. Gastropoda: Prosobranchia: Hydro- 
biidae. | 

cahuillarum, Pyrgulopsis — Taylor, 1950: 31— 
32, fig. 7. Type locality: Lake Cahuilla, 50 
yards NE of the so-called Fish Traps, 7.9 mi 
W of Mecca, 33°34’17”М, 116°13’13”W, Riv- 
erside Co., California. Late Pleistocene? 
Holotype: SBMNH 35503 (ex S. S. Berry 
collection 13257). Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Taylor (1966c: 
196) subsequently concluded that this was 
a junior synonym of Tryonia protea (Gould, 
1855): 

Calibasis Taylor, 1966b: 41. Described as a 
subgenus of Juga H. Adams & A. Adams, 
1854. Type species Juga acutifilosa 
(Stearns, 1890); original designation. Gas- 
tropoda: Prosobranchia: Pleuroceridae. 

californica, Batillaria — Taylor, 1983: 290, fig. 
1. Type locality: USGS locality M3091, 5,600 
ft W, 2,800 ft N, in irregular sec. 20, T. 10S, 
К. 21E, Imperial Co., California. Bouse For- 
mation, Upper Miocene - Pliocene. Holotype 
USNM 305208; paratypes USNM 305209- 
305211. Gastropoda: Prosobranchia: 
Potamididae. 

californiensis, Fontelicella (Fontelicella) — 
Gregg & Taylor, 1965: 109. Type locality, 
Campo Creek, 0.6 mi E of Mountain Empire 
Dam, W 1/2 SW 1/4 sec. 19, T. 18S, R. 5E, 
San Diego Co., California. Holotype, UMMZ 
220000 (only specimen). Gastropoda: 
Prosobranchia: Hydrobiidae. Notes: Hershler 
(1994: 25) recognized this taxon as a valid 
species of Pyrgulopsis Call & Pilsbry, 1886. 

caramba, Paludiscala — Taylor, 1966c: 207- 
208, pl. 13, figs. 11, 14, 16, text-figs. 23-25. 
Type locality: Spring tributary to the area of 
marshes and lagunas named “El Mojarral,” 
1.7 km due E of the northern tip of Sierra de 
San Marcos, 11 km SW of Cuatro Ciénegas, 
Coahuila, Mexico. Holotype, UMMZ 220164. 
Gastropoda: Prosobranchia: Hydrobiidae. 
Notes: Hershler (1985: 60-64) redescribed 
this taxon. 


Caribnauta Taylor, 2003: 47-48. Type species 
Caribnauta harryi Taylor, 2003; original des- 
ignation. Gastropoda: Pulmonata: Physidae. 
Notes: Taylor (2002a: 25) previously used 
this name as a nomen nudum. 

carinifex, Menetus — Taylor in Taylor & Smith, 
1981: 366-367, pls. 11-12. Type locality: 
Mopung Hills, in coquina and limestone 
beds, between Southern Pacific Railroad 
tracks and the Fallon-Lovelock Road, mostly 
in the NW 1/4 sec. 7, T. 23N, R. 29E, 
Churchill Co., Nevada. Pliocene. Holotype 
UMMZ 250115. Gastropoda: Pulmonata: 
Planorbidae. 

carranzae, Mexipyrgus — Taylor, 1966c: 190, 
pl. 15, figs. 27-32, text-fig. 16. Type locality: 
Laguna Tio Candido, 14 km S of Cuatro 
Ciénegas, Coahuila, Mexico. Holotype 
UMMZ 220211. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: Hershler (1985: 87) ini- 
tially treated this species as a junior synonym 
of Mexipyrgus churinceanus Taylor, 1966, 
but as M. carranzae was the type species of 
Mexipyrgus Taylor, 1966, Hershler & Thomp- 
son (1982: 78) restored M. carranzae as a 
valid species, with M. churinceanus and four 
other species as junior synonyms. 

chauliodonta, Gastrocopta (Gastrocopta) — 
Taylor, 1954d: 12. Type locality: Center of S 
side of NW 1/4 section 25, T. 31N, К. 22W, 
Brown Co., Nebraska. Sand Draw fauna, late 
Nebraskan age, Pleistocene. Holotype 
UMMZ 181120; paratypes UMMZ 181121. 
Gastropoda: Pulmonata: Pupillidae. 

Chiapaphysa Taylor, 2003: 167. Type species 
Chiapaphysa grijalvae Taylor, 2003; original 
designation. Gastropoda: Pulmonata: 
Physidae. Notes: Taylor (2002a: 25) previ- 
ously used this name as a nomen nudum 
(as “Chipaphysa” [sic]). 

chippevarum, Laurentiphysa — Taylor, 2003: 
154-157, figs. 148-152, pl. 8, fig. 1, map 
fig. 147. Type locality: Ditch on N side of 
Highway 77, 1.85 mi W of Highway 13, SW 
1/4 SE 1/4 sec. 31, T. 43N, R. 2W, elevation 
1,530 ft, Ashland Co., Wisconsin. Holotype 
CAS 146089; paratypes CAS 146090; 
BMNH 20001309. Gastropoda: Pulmonata: 
Physidae. 

chrysopylica, Juga — Taylor, 1966b: 39. For 
“Goniobasis rodeoensis” sensu Hanna 
(1923: 34-35, pl. 1, fig. 3), non В; L. Clark 
(1915: 491-492, pl. 69, figs. 1, 10, as 
Cerithium rodeoensis). Sonoma Co., Cali- 
fornia. Petaluma Formation, Pliocene. Gas- 
tropoda: Prosobranchia: Pleuroceridae. 


DWIGHT WILLARD TAYLOR 203 


Notes: The syntypic type material would be 
Hanna’s figured specimens, CAS 70423- 
70428. 

chupaderae, Fontelicella — Taylor, 1987: 24- 
26, fig. 11. Type locality Willow Spring, on 
Cienaga Ranch at S end of Chupadera 
Mountains, about 5 mi W of Bosque del 
Apache National Wildlife Refuge headquar- 
ters, Socorro Co., New Mexico. Holotype 
LACM 2218; paratypes UTEP 10052; ANSP 
376027; FLMNH 160938; USNM 854081. 
Gastropoda: Prosobranchia: Hydrobiidae. 
Notes: Hershler (1994: 30) recognized this 
taxon as a valid species of Pyrgulopsis Call 
& Pilsbry, 1886. 

churinceanus, Mexipyrgus — Taylor, 1966c: 
190-201, pl. 16, figs. 33-39, pl. 17, figs. 40, 
41. Type locality: Laguna Churince, 16 km 
SW of Cuatro Ciénegas, Coahuila, Mexico. 
Holotype UMMZ 220150. Gastropoda: 
Prosobranchia: Hydrobiidae. Notes: Hershler 
(1985: 87-104) and Hershler & Thompson 
(1992: 78) concluded that this species was 
a junior synonym of Mexipyrgus carranzae 
Taylor, 1966. 

Clenchiellini Taylor, 1966c: 175, 181. New tribe 
within the Cochliopinae for Clenchiella 
Abbott, 1948. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: Subsequently elevated 
to a subfamily by Starobogatov (1970: 34), 
and then to a full family by loganzen & 
Starobogatov (1982: 1114). Ponder & 
Bouchet (2005: 252) listed this as a subfam- 
ily of the Hydrobiidae sensu stricto. 
coahuilae, Durangonella — Taylor, 1966c: 184— 
186, pl. 14, figs. 19, 22. Type locality: La- 
guna Grande, in the middle of the E side, 
within about 30 m of the mouth of Rio 
Churince and about 17 km SW of Cuatro 
Ciénegas, Coahuila, Mexico. Holotype 
UMMZ 220159. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: Hershler et al. (2002: 
172-182) concluded that Durangonella 
Morrison, 1945, was a junior synonym of 
Tryonia Stimpson, 1865, and described 
Juturnia as a new genus for this species (as 
the type species) and two other species. 
Coahuilix Taylor, 1966c: 180. Type species 
Coahuilix hubbsi Taylor, 1966; original des- 
ignation. Taylor classified this in the tribe 
Horatiini, subfamily Cochliopinae. Gas- 
tropoda: Prosobranchia: Hydrobiidae. Notes: 
Hershler (1985: 53-54) redescribed this 
taxon. 

“Cochliopinae” Taylor, 1966c: 173. Although 
Taylor listed this as a new subfamily, this 


taxon was already established by Tryon 
(1866) as a subfamily in the Amnicolidae 
[Hydrobiidae]. 

“Cochliopini” Taylor, 1966c: 173. Although Tay- 
lor listed this as a new tribe, this taxon was 
first established by Tryon (1866) as a sub- 
family in the Amnicolidae [Hydrobiidae], so 
Taylor is not to be credited as the author of 
this tribe, pursuant to ICZN Article 36.1 (“A 
name established for a taxon at any rank in 
the family group is deemed to have been 
simultaneously established for nominal taxa 
at all other ranks in the family group ... The 
name has the same authorship and date at 
every rank.”). 

conchos, Disconaias — Taylor, 1997b: 420- 
423, pl. 21, fig. 1. Type locality: Rio Conchos, 
about 0.5 km W of Julimes, Chihuahua, 
Mexico. Holotype LACM 2257; paratype 
MCZ 316166. Bivalvia: Unionidae. 

Craterarion Taylor, 1954c: 75. Type species 
Craterarion pachyostracon Taylor, 1954: 
Original designation. Upper Miocene. Gas- 
tropoda: Pulmonata: Arionidae. 

davisi, Fontelicella — Taylor, 1987: 10-12, fig. 
4. Type locality: tributary of Limpia Creek, 
about 5 mi NE of Fort Davis, Jeff Davis Co., 
Texas. Holotype LACM 2211; paratypes 
UTEP: 10053; ANSP: 376023; FLMNE 
160944; USNM 854085. Gastropoda: 
Prosobranchia: Hydrobiidae. Notes: Hershler 
(1994: 32) recognized this taxon as a valid 
species of Pyrgulopsis Call & Pilsbry, 1886. 

dineana, Limnaea (Pseudosuccinea) — Taylor, 
1957b: 659-660, figs. 4-6. Type locality: 
White Cone Peak, sec. 12, T. 25N, R. 21E, 
1.5 mi S of White Cone Trading Post, Na- 
vajo Co., Arizona. White Cone local fauna, 
Bidahochi Formation, Hemphillian Age, 
Middle Pliocene. Holotype USNM 562084; 
paratypes USNM 562086. Gastropoda: 
Pulmonata: Lymnaeidae. Notes: Taylor in 
Taylor & Smith (1981: 363) transferred this 
species to Lutrilimnea. 

escobedae, Mexipyrgus — Taylor, 1966c: 191- 
192, pl. 14, figs. 23-26. Type locality: La- 
guna Escobeda, 12 km $ of Cuatro 
Ciénegas, Coahuila, Mexico. Holotype 
UMMZ 220202. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: Hershler (1985: 87-104) 
and Hershler & Thompson (1992: 78) con- 
cluded that this species was a junior syn- 
onym of Mexipyrgus carranzae Taylor, 1966. 

Fontelicella Gregg & Taylor, 1965: 103-108. 
Type species Fontelicella californiensis 
Gregg & Taylor, 1965; original designation. 


204 KABAT & JOHNSON 


Gastropoda: Prosobranchia: Hydrobiidae. 
Notes: Thompson (1979: 47) classified 
Fontelicella in the Nymphophilinae; subse- 
quently, Hershler & Thompson (1987: 28) 
and Hershler (1994: 5-11) concluded that 
Fontelicella was a junior synonym of 
Pyrgulopsis Call & Pilsbry, 1886. 

Fontigentinae Taylor, 1966c: 182. New sub- 
family for Fontigens Pilsbry 1933. Gas- 
tropoda: Prosobranchia: Hydrobiidae. Notes: 
Hershler et al. (1990) synonymized Fonti- 
gentinae with Emmericiinae Brusina, 1870, 
which is where Morrison (1949) had previ- 
ously classified Fontigens. Ponder & Bouchet 
(2005: 251) classified the Emmericiinae in 
the family Amnicolidae. 

franzenae, Gastrocopta — Taylor, 1960b: 67- 
69, pl. 1, fig. 29. Type locality: Rexroad 
Ranch, University of Kansas Meade Co. loc. 
3, USGS Cenozoic loc. 21171, W 1/2 SW 1/ 
4 sec. 22, T. 33$, К. 29W, Kansas. Upper 
Pliocene. Holotype UMMZ 183033-a. Gas- 
tropoda: Pulmonata: Pupillidae. 

gentilis, Lutrilimnea — Taylor in Taylor & Smith, 
1981: 363-364, pl. 8. Type locality: SE 1/4 
SW 1/4 sec. 12, T. 115, В. 40E, Lake 
Thatcher, Caribou Co., Idaho. Main Canyon 
Formation, Pleistocene. Holotype UMMZ 
250110. Gastropoda: Pulmonata: Lymnaei- 
dae. 

gilae, Fontelicella — Taylor, 1987: 16-18, fig. 
7. Type locality: springs on N side of East 
Fork of Gila River, center of sec. 3, T. 13$, 
R. 13W, Grant Co., New Mexico. Holotype 
LACM 2214; paratypes UTEP 10054; ANSP 
376025; FLMNH 160936; USNM 854087. 
Gastropoda: Prosobranchia: Hydrobiidae. 
Notes: Hershler (1994: 36) recognized this 
taxon as a valid species of Pyrgulopsis Call 
& Pilsbry, 1886. 

gilae, Тгуота — Taylor, 1987: 36-37, fig. 17. 
Type locality: unnamed spring on N side of 
Gila River, about 2 mi N of Bylas, in T. 3S, R. 
22E, 25,000 ft W and 15,000 ft N of the town- 
ship line, Graham Co., Arizona. Holotype 
LACM 2187; paratypes UTEP 10063; ANSP 
376029; FLMNH 160943, 160948; USNM 
854074, 854089. Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Tryonia gilae 
Hershler in Hershler & Landye, 1988: 43-49 
is a junior subjective synonym and junior pri- 
mary homonym of Tryonia gilae Taylor, 1987 
(fide Hershler & Landye 1988: 58). Hershler 
(2001: 9-10) redescribed this species. 

grijalvae, Chiapaphysa — Taylor, 2003: 168- 
170, figs. 166, 167, pl. 8, fig. 7, map fig. 165. 
Type locality: Rio Suchiapa, 2 km SE of 


Suchiapa, 16°36.4’N, 93°5.0’W, Chiapas, 
Mexico. Holotype CAS 114818; paratypes 
CAS 114787; BMNH 20001308; MCZ 
302595; ZIBM CNMO 1161. Gastropoda: 
Pulmonata: Physidae. 


harpa, Physa (Costatella) — Taylor in Taylor & 


Smith, 1981: 368-370, pl. 16. Type locality 
USGS loc. 20093, SE 1/4 sec. 16, T. 13N, 
R. 2W, Box Elder Co., Utah. Cache Valley 
Formation, Pliocene. Holotype USNM 
305783; paratypes USNM 781782, 781783. 
Gastropoda: Pulmonata: Physidae. 


harryi, Caribnauta — Taylor, 2003: 48-49, fig. 


20, pl. 1, fig. 2, map fig. 5. Type locality: 
stream W of Las Piedras, Puerto Rico. For 
“Physa marmorata” sensu Harry & 
Hubendick, 1964, non Guilding, 1828. Gas- 
tropoda: Pulmonata: Physidae. Notes: Tay- 
lor wrote that the holotype was the “shell 
figured by Harry & Hubendick (1964, fig. 72),” 
and that “| wrote to the [Göteborg] Museum 
for information. No reply was received.” Tay- 
lor also wrote that: “Paratypes might be in 
the Houston Museum of Natural Science, 
Houston, Texas, where Harry's collection is 
deposited. | wrote to the museum asking for 
loan of the relevant specimens, but received 
the reply that other duties prevented the cu- 
rator from aiding me.” 


Haitini Taylor, 2003: 128. New tribe in subfam- 


ily Physinae, for Haitia Clench & Aguayo, 
1932. Gastropoda: Pulmonata: Physidae. 
Notes: Taylor (2002a: 25) previously used 
this name as a nomen nudum. 


hemphilli, Physella — Taylor, 2003: 191, pl. 9, 


fig. 5, map fig. 8. Type locality: Lake Coeur 
d’Alene, Kootenai Co., Idaho. Holotype CAS 
116331; paratypes CAS 114824, CAS 
114825. Gastropoda: Pulmonata: Physidae. 
Notes: Taylor (2002a: 25) previously used 
this name as a nomen nudum. 


Horatiini Taylor, 1966c: 179-180. New tribe in 


subfamily Cochliopinae, for Horatia 
Bourguignat, 1887, and eleven other gen- 
era: Coahuilix, Gocea, Hadziella, Daude- 
bardiaella, Hauffenia, Neohoratia, Lyhnidia, 
Ohridohoratia, Ohrigocea, and Karevia. Gas- 
tropoda: Prosobranchia: Hydrobiidae. Notes: 
Hershler & Thompson (1992: 129) rejected 
Taylor s combination of genera from south- 
eastern Europe and northern Mexico into a 
single tribe. Ponder & Bouchet (2005: 252) 
listed Horatiini as a junior synonym of 
Belgrandiinae de Stefani, 1877, in the 
Hydrobiidae sensu stricto; Wilke et al. (2001: 
21) listed Horatiinae as a valid subfamily in 
the Hydrobiidae sensu Stricto. 


DWIGHT WILLARD TAYLOR 205 


hubbsi, Coahuilix — Taylor, 1966c: 180-181, 
figs. 8-13. Type locality: from a bottom 
sample of the northernmost pool of Pozos 
de la Becerra, 14 km SW of Cuatro Ciéne- 
gas, Coahuila, Mexico. Holotype UMMZ 
220180. Gastropoda: Prosobranchia: Hydro- 
biidae. Notes: Hershler (1985: 54-57) re- 
described this species. 

humboldtina, Physa (Costatella) — Taylor in 
Taylor & Smith, 1981: 367-368, pls. 13-14. 
Type locality: Mopung Hills, in coquina and 
limestone beds, between Southern Pacific 
Railroad tracks and the Fallon-Lovelock 
Road, mostly in the NW 1/4 sec. 7, T. 23N, 
R. 29E, Churchill Co., Nevada. Pliocene. 
Holotype UMMZ 250118. Gastropoda: 
Pulmonata: Physidae. 

Idabasis Taylor, 1966b: 41. Described as a 
subgenus of Juga H. Adams & A. Adams, 
1854. Type species Juga chrysopylica Tay- 
lor, 1966; original designation. Pliocene. 
Gastropoda: Prosobranchia: Pleuroceridae. 

idahoensis, Valvata — Taylor in Taylor & Smith, 
1981: 358. Replacement name for Valvata 
multicarinata Yen, 1946 (pp. 487-488, pl. 76, 
fig. 1), non Hislop, 1860 (pp. 170-171, pl. 5, 
fig. 15a, b). Bear Lake Co., Idaho. Salt Lake 
Group, Pliocene. Holotype USNM 559943; 
paratype USNM 559944. Gastropoda: 
Heterobranchia: Valvatidae. 

imminens, Pyrgulopsis — Taylor, 1950: 28, figs. 
1-3. Type locality: shore of Salton Sea, by 
Fish Springs, Imperial Co., California. Holo- 
type SBMNH 35497 (ex S. S. Berry coll. 
13255); paratypes SBMNH 35498 and 35499 
(ex Sx So Beny colli 13252 and 13283); 
SDSNH; USNM 613967; UMMZ. Gas- 
tropoda: Prosobranchia: Hydrobiidae. Notes: 
Taylor stated that another paratype was in 
the S. S. Berry collection, No. 13258, but that 
specimen could not be found at SBMNH. 
Hershler & Thompson (1992: 111) concluded 
that this taxon was a junior synonym of 
Tryonia protea (Gould, 1855). 

intermontana, Radix — Taylor, 1966b: 68. Re- 
placement name for Lymnaea idahoensis 
Yen (1946: 490, pl. 76, fig. 8), non Hender- 
son, 1931. Bear Lake Co., Idaho. Salt Lake 
Group, Pliocene. Holotype USNM 559955: 
paratype USNM 559956. Gastropoda: 
Pulmonata: Lymnaeidae. 

jaliscoensis, Amecanauta — Taylor, 2003: 73- 
74, figs. 49-52, pl. 1, fig. 3, map fig. 15. Type 
locality: roadside ditch on W side of Mexico 
200, opposite entrance to “Modulo de Abasto” 
de Puerto Vallarta, 2.2 km NE of entrance to 
airport, 20°41.48’N, 105°13.95’W, Jalisco, 


Puerto Vallarta. Holotype CAS 114813; 
paratypes CAS 114800; BMNH 20001306; 
MCZ 302596; ZIBM CNMO 1159. Gas- 
tropoda: Pulmonata: Physidae. 

junturae, Fluminicola — Taylor, 1963a: 38, figs. 
8-10. Type locality: USGS loc. 21173, 
Malheur Co., Oregon. Black Butte local 
fauna, Juntura Formation, Miocene. Holotype 
USNM 563115. Gastropoda: Prosobranchia: 
Hydrobiidae. Transferred to Lithoglyphus by 
Taylor (1975: 105). 

junturae, Radix — Taylor, 1963a: 38, 40, figs. 
11-18. Type locality: USGS loc. 21173, 
Malheur Co., Oregon. Black Butte local 
fauna, Juntura Formation, Miocene. Holotype 
USNM 563107; paratypes USNM 563108, 
563109. Gastropoda: Prosobranchia: 
Lymnaeidae. 

kolobensis, Fontelicella — Taylor, 1987: 19-20, 
fig. 8. Type locality: Toquerville Springs, sec. 
35, T. 40S, R. 13W, Washington Co., Utah. 
Holotype LACM 2216; FLMNH 160940; 
USNM 854076. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: Hershler (1994: 44) rec- 
ognized this taxon as a valid species of 
Pyrgulopsis Call & Pilsbry, 1886, and con- 
cluded that Fontelicella pinetorum Taylor, 
1987, was a junior synonym of this taxon. 

kosteri, Tryonia (Paupertryonia) — Taylor, 1987: 
45-47, fig. 23. Type locality: Sago Spring, 
900 ft W, 2,400 ft S, sec. 5, T. 10S, R. 25E, 
Chaves Co., New Mexico. Holotype LACM 
2252; paratypes UTEP 10064; ANSP 
376028; FLMNH 160950, 160951; USNM 
854081, 854091. Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Hershler 
(2001: 15) redescribed this species and 
transferred it to Durangonella Morrison, 
1945; subsequently, Hershler et al. (2002: 
172-182) concluded that Durangonella was 
a junior synonym of Tryonia Stimpson, 1865, 
and described Juturnia as a new genus for 
this and two other species. 

lasseni, Vorticifex — Taylor in Taylor & Smith, 
1981: 349, 352-353, pl. 2, figs. 8, 9. Type 
locality: shore of historical Honey Lake (now 
dry), at about 3,980 ft, Lassen Co., Califor- 
nia. Holotype UMMZ 250100. Gastropoda: 
Pulmonata: Planorbidae. 

Laurentiphysa Taylor, 2003: 152. Type spe- 
cies: Physa vernalis Taylor & Jokinen, 1985; 
Original designation. Gastropoda: Pulmo- 
nata: Physidae. Notes: Taylor (2002a: 25) 
previously used this name as a nomen nu- 
dum. 

lavernensis, Gastrocopta (Gastrocopta) — Tay- 
lor, 1954d: 11. Type locality: NW 1/4 sec. 5, 


206 KABAT & JOHNSON 


T. 3N, R. 28E, Beaver Co., Oklahoma. 
Laverne Formation, Lower Pliocene. Holo- 
type UMMZ 181274; paratypes UMMZ 
181275. Gastropoda: Pulmonata: Pupillidae. 

“Lithoglyphinae” Taylor, 1966c: 182. As a new 
subfamily for Lithoglyphus Hartman, 1821, 
but this taxon was already established as a 
family-level taxon by Tryon (1866: 156), so 
Taylor is not to be credited as the author of 
this subfamily, pursuant to ICZN Article 36.1 
(“A name established for a taxon at any rank 
in the family group is deemed to have been 
simultaneously established for nominal taxa 
at all other ranks in the family group ... The 
name has the same authorship and date at 
every rank.”). Gastropoda: Prosobranchia: 
Hydrobiidae. 

“Littoridininae” Taylor, 1966c: 182. As a new 
subfamily for Littoridina and numerous gen- 
era, but this taxon was already established 
as a family-level taxon by Thiele (1928: 372, 
378), so Taylor is not to be credited as the 
author of this subfamily, pursuant to ICZN 
Article 36.1 (“A name established for a taxon 
at any rank in the family group is deemed to 
have been simultaneously established for 
nominal taxa at all other ranks in the family 
group ... The name has the same author- 
ship and date at every rank.”). Gastropoda: 
Prosobranchia: Hydrobiidae. 

lugoi, Mexipyrgus — Taylor, 1966c: 192, pl. 17, 
figs. 42—45. Type locality: Rio Mesquites, at 
the main road, 9 km SW of Cuatro Ciénegas, 
Coahuila, Mexico. Holotype UMMZ 220185. 
Gastropoda: Prosobranchia: Hydrobiidae. 
Notes: Hershler (1985: 87-104) and 
Hershler & Thompson (1992: 78) concluded 
that this species was a junior synonym of 
Mexipyrgus carranzae Taylor, 1966. 

Lutrilimnea Taylor in Taylor & Smith, 1981: 360— 
361. Type species Lutrilimnea polyskelidis 
Taylor in Taylor & Smith, 1981; original des- 
ignation. Gastropoda: Pulmonata: Lymna- 
eidae. 

Mayabina Taylor, 2003: 88-92. Type species 
Physa cisternina Morelet, 1851; original des- 
ignation. Gastropoda: Pulmonata: Physidae. 
Notes: Taylor (2002a: 25) previously used 
this name as a nomen nudum. Taylor (2003: 
88, 104) stated that Physa cisternina 
Morelet, 1851, was the type species, but he 
also listed that taxon as a junior synonym of 
Physa spiculata Morelet, 1849. 

megachlamys [sic], Physa — Taylor in Wu & 
Brandauer, 1982: 7, nomen nudum. 

megalochlamys, Physa — Taylor, 1988c: 55- 
62, fig. 3. Type locality: Lily Pond besides 


US Highway 26-89-187, NW 1/4 sec. 19, T. 
45N, R. 114W, Teton Co., Wyoming. Holo- 
type LACM 2255; paratypes UC 30260. Gas- 
tropoda: Pulmonata: Physidae. Notes: 
Although Taylor stated that the holotype was 
deposited in the LACM, it cannot now be 
found there and evidently was never depos- 
ited or was later removed from the incipient 
type collection. Taylor (2003: 164, figs. 159- 
162, pl. 8, fig. 3) redescribed this taxon, and 
stated that the holotype was “CAS 114779,” 
presumably the specimen that was sup- 
posed to have been deposited in the LACM. 

melina, Fontelicella (Natricola) — Taylor in Tay- 
lor & Smith, 1981: 348, 350-352, pl. 2, figs. 
1, 2. Type locality: Honey Lake, SE 1/4 NW 
1/4 sec. 27, elevation 3,995 ft, Lassen Co., 
California. Pleistocene? Holotype UMMZ 
250095. Gastropoda: Prosobranchia: Hydro- 
biidae. 

metcalfi, Fontelicella — Taylor, 1987: 12-14, 
fig. 5. Type locality: Naegele Springs, 5.3 mi 
NNW of Ruidosa, Presidio Co., Texas. Ho- 
lotype LACM 2212; paratypes UTEP 10055; 
ANSP 376024; FLMNH 160937; USNM 
854077. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: Hershler (1994: 49) rec- 
ognized this taxon as a valid species of 
Pyrgulopsis Call & Pilsbry, 1886. 

Mexinauta Taylor, 2003: 74-76. Type species 
Physa nitens Philippi, 1841; original desig- 
nation. Gastropoda: Pulmonata: Physidae. 
Notes: Taylor (2002a: 25) previously used 
this name as a nomen nudum. 

Mexipyrgus Taylor, 1966c: 188-189. Type spe- 
cies Mexipyrgus carranzae Taylor, 1966; 
original designation. Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Hershler 
(1985: 87-105) reviewed the six included 
species and concluded that they all repre- 
sented a single polytypic taxon. Hershler & 
Thompson (1992: 75-78) transferred this 
genus from the Littoridininae to the Coch- 
liopinae. 

Mexithauma Taylor, 1966c: 205. Type species 
Mexithauma quadripaludium Taylor, 1966; 
original designation. Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Hershler 
(1985: 72-78) transferred this monotypic 
genus from Mexithaumatinae to Littoridininae 
(= Cochliopinae) and redescribed the type 
species. 

Mexithaumatinae Taylor, 1966c: 204. De- 
scribed as a subfamily, questionably placed 
in the Hydrobiidae, for Mexithauma Taylor, 
1966. Gastropoda: Prosobranchia: Hydro- 
biidae. Notes: Hershler (1984: 65-66) and 


DWIGHT WILLARD TAYLOR 207 


Hershler & Thompson (1992: 6) concluded 
that this subfamily was a junior synonym of 
Cochliopinae Tryon, 1866 (formerly Litto- 
ridininae). 

Microamnicola Gregg & Taylor, 1965: 109. De- 
scribed as a subgenus of Fontelicella Gregg 
& Taylor, 1965. Type species Fontelicella 
(Microamnicola) micrococcus (Pilsbry in 
Stearns, 1893); original designation. Gas- 
tropoda: Prosobranchia: Hydrobiidae. Notes: 
Hershler & Thompson (1987: 28) and 
Hershler (1994: 5-11) concluded that this 
taxon was a junior synonym of Pyrgulopsis 
Call & Pilsbry, 1886. 

micromphalus, Menetus (?) — Taylor, 1954c: 
74—75, pl. 20, figs. 4-9. Type locality: W end 
of Barstow Hills, 7 mi N of Barstow, “Lake 
Bed Horizon” in the canyon next S of Pirie 
Canyon, middle of SW 1/4, sec. 15, T. 11N, 
R. 2W, San Bernardino Co., California. 
Barstow Formation, Upper Miocene. Holo- 
type Taylor collection 2038a; paratypes Tay- 
lor collection 2038; CAS 70416 (ex SU 
8079), CAS 70417 (ex SU 8080); USNM 
561451. Gastropoda: Pulmonata: Planorbi- 
dae. 

milleri, Cochliopina — Taylor, 1966c: 177-178, 
text figs. 6, 7, pl. 13, figs. 12, 13. Type local- 
ity: Rio Mesquites, at the main road 9 km 
SW of Cuatro Ciénegas, Coahuila, Mexico. 
Holotype UMMZ 220182. Gastropoda: 
Prosobranchia: Hydrobiidae. Notes: Hershler 
(1985: 68-70) redescribed this taxon. 

minckleyi, Nymphophilus — Taylor, 1966c: 199- 
203, pl. 13, figs. 15, 17, text figs. 17-21. Type 
locality: Rio Mesquites, at the main road 9 
km SW of Cuatro Ciénegas, Coahuila, 
Mexico. Holotype UMMZ 220188. Gas- 
tropoda: Prosobranchia: Hydrobiidae. Notes: 
Hershler (1985: 38-45) redescribed this 
taxon. 

minutus, Promenetus — Taylor, 1954a: 37-38. 
Type locality: Allee Stream, opposite labo- 
ratory, Barro Colorado Island, Gatun Lake, 
Panama Canal Zone, Panama. Holotype 
USNM 605858 (ex USNM 588913): 
paratypes USNM 588913; UMMZ 181122. 
Gastropoda: Pulmonata: Planorbidae. 

mirolli, Physa — Taylor, 2003: 165-166. For 
Physa fontinalis sensu Mirolli (1958: 245- 
247, pls. 24-26) non Physa fontinalis 
(Linnaeus, 1758). Italy. Holotype “the shell 
illustrated by Mirolli (1958: pl. 24, fig. 1).” 
Gastropoda: Pulmonata: Physidae. Notes: 
Taylor (2002a: 25) previously used this name 
as a nomen nudum. 


mohaveana, Lymnaea (Stagnicola) — Taylor, 
1954c: 73, pl. 20, figs. 1, 2. Type locality: W 
end of Barstow Hills, 7 mi N of Barstow, 
middle of SE 1/4 sec. 15, T. 11N, R. 2W., 
San Bernardino Co., California. Barstow 
Formation, Upper Miocene. Holotype SU 
8077 (not located in the CAS, Jan. 2008): 
paratypes CAS 70415 (ex SU 8078); Taylor 
coll. 1785, 2035; USNM 561488; UCMP 
34194-34197. Gastropoda: Pulmonata: 
Lymnaeidae. 

mojarralis, Mexipyrgus — Taylor, 1966c: 193, 
pl. 18, figs. 46-49, 51-53. West Laguna in 
El Mojarral, 1.7 km ENE of the northern tip 
of Sierra de San Marcos, Coahuila, Mexico. 
Holotype UMMZ 220192. Gastropoda: 
Prosobranchia: Hydrobiidae. Notes: Hershler 
(1985: 87-104) and Hershler & Thompson 
(1992: 78) concluded that this species was 
a junior synonym of Mexipyrgus carranzae 
Taylor, 1966. 

moreleti, Haitia — Taylor, 2003: 146-147, fig. 
140, pl. 6, fig. 2, map fig. 15. Type locality: 
marshy border of Lake Peten-Itza, Santa 
Elena, El Petén, Guatemala. Holotype CAS 
114821. Gastropoda: Pulmonata: Physidae. 

multilineatus, Mexipyrgus — Taylor, 1966c: 
193-194, pl. 18, figs. 50, 54-57. Type local- 
ity: East Laguna in El Mojarral, 1.9 km ENE 
of the northern tip of Sierra de San Marcos, 
Coahuila, Mexico. Holotype UMMZ 220197. 
Gastropoda: Prosobranchia: Hydrobiidae. 
Notes: Hershler (1985: 87-104) and 
Hershler & Thompson (1992: 78) concluded 
that this species was a junior synonym of 
Mexipyrgus carranzae Taylor, 1966. 

Natricola Gregg & Taylor, 1965: 108-109. 
Described as a subgenus of Fontelicella 
Gregg & Taylor, 1965. Type species Fonteli- 
cella (Natricola) robusta (Walker, 1908); 
original designation. Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Hershler & 
Thompson (1987: 28) and Hershler (1994: 
9-11) concluded that this taxon was a junior 
synonym of Pyrgulopsis Call & Pilsbry, 1886. 
Hershler & Liu (2004a) discussed the three 
Recent species included by Gregg & Taylor 
in “Natricola” and concluded that only the 
type species was valid, with the others be- 
ing junior synonyms. 

natricina, Physa (Haitia) — Taylor, 1988c: 67, 
fig. 6, map fig. 7. Type locality: Snake River, 
in rapids along E side, SW 1/4 SE 1/4 sec. 
21, Т. 6S, К. 13E., Gooding Co., Idaho. Ho- 
lotype LACM “2256” [= LACM 2970]. Gas- 
tropoda: Pulmonata: Physidae. Notes: As 


208 


LACM 2256 was inadvertently used for an- 
other author’s type specimen, Taylor’s holo- 
type was catalogued as LACM 2970. Taylor 
(2003: 147) erroneously stated that the ho- 
lotype was “CAS 114795.” Rogers & 
Wethington (2007) discussed the “paratypes” 
of this species, but Taylor only specified the 
holotype, and the remaining listed specimens 
were not given any type status, so there are 
no paratypes (ICZN Articles 72.4.6 and 
73.1.1). They also stated that they could not 
find the “paratypes” in the “US Geological 
Survey Western Ecological Resource Cen- 
ter,” but the USGS numbers in Taylor’s de- 
scription are USGS station numbers, not 
specimen catalog numbers. Rogers & 
Wethington (2007) concluded that this taxon 
was a junior synonym of Physa acuta 
Draparnaud, 1805; these authors are con- 
tinuing their genetic studies of these and 
related taxa (A. Wethington to A. R. Kabat, 
in litt., Nov. 29, 2007). 

nevadense, Sphaerium (Amesoda) — Taylor in 
Taylor & Smith, 1981: 356357, pl. 3, figs. 1— 
6. Type locality: limestone and coquina beds, 
NW 1/4 sec. 17, T. 23N, R. 29E, Mopung 
Hills, Churchill Co., Nevada. Pliocene. Ho- 
lotype UMMZ 250102 [right valve only]. 
Bivalvia: Sphaeriidae. 

nevadensis, Valvata — Taylor in Taylor & Smith, 
1981: 357-358, pl. 3, figs. 7-12. Type local- 
ity: limestone and coquina beds, SW 1/4 sec. 
17, T. 23N, R. 29E, strike N 60° W, dip 4° 
NE, thickness 15-20 ft, weathering on the 
service 1100-1300 ft E, 1300-1400 ft N of 
the SW section corner, Mopung Hills, 
Churchill Co., Nevada. Pliocene. Holotype 
UMMZ 250104. Gastropoda: Heterobran- 
chia: Valvatidae. 

Nymphophilinae Taylor, 1966c: 199. Described 
as a subfamily in the Hydrobiidae, for 
Nymphophilus Taylor, 1966. Gastropoda: 
Prosobranchia: Hydrobiidae. Notes: Thomp- 
son (1979) redescribed this subfamily, which 
he treated as a senior synonym of Ori- 
entaliidae Radoman, 1973, and concluded 
that 14 other genera should be included. 
Hershler et al. (2003) concluded that the 
North American taxa of this subfamily prob- 
ably formed a monophyletic clade. Ponder 
& Bouchet (2005: 252) listed Nym- 
phophilinae as a valid subfamily in the 
Hydrobiidae s. str. 

Nymphophilus Taylor, 1966c: 199. Type spe- 
cies Nymphophilus minckleyi Taylor, 1966; 
original designation. Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Thompson 


KABAT & JOHNSON 


(1979) and Hershler (1985: 38-45) re- 
described this genus and its included spe- 
cies. Liu & Hershler (2005: 291-292, 296) 
concluded that Nymphophilus “is little differ- 
entiated from Pyrgulopsis apart from the 
unusual shell of its type species,” and the 
cladistic analyses consistently placed the 
species of the former genus within the latter 
genus, thereby warranting treating Nympho- 
philus as a junior synonym of Pyrgulopsis 
Call & Pilsbry, 1886. 


Oreobasis Taylor, 1966b: 41. Described as a 


subgenus of Juga H. Adams & A. Adams, 
1854. Type species Melania newberryi Lea, 
1860; original designation. Taylor also sug- 
gested that this species was “probably” a 
junior synonym of Melania bulbosa Gould, 
1847. Gastropoda: Prosobranchia: Pleuro- 
ceridae. 


Oreoconus Taylor in McKenna et al., 1962: 11. 


Type species Oreoconus planispira Taylor in 
McKenna et al., 1962; original designation. 
Gastropoda: Pulmonata: Bulimulidae. 


pachyostracon, Craterarion — Taylor, 1954c: 


75, pl. 20, figs. 16-20. Type locality: W end 
of Barstow Hills, 7 mi N of Barstow, “Lake 
Bed Horizon” in the canyon next S of Pirie 
Canyon, middle of SE 1/4, sec. 15, T. 11N, 
R. 2W, San Bernardino Co., California. 
Barstow Formation, Upper Miocene. Holo- 
type CAS 70410 (ex Stanford Univ. 8073); 
paratypes CAS 70412 (ex Stanford Univ. 
8074), 70411 (ex CAS 10215, 10215a-k); 
Taylor collection 1791; $. $. Berry collection 
19936; LACMIP 4920, 4921 (ex W.O. Gregg 
collection 5924); UCMP 34183; USNM 
561449. Gastropoda: Pulmonata: Arionidae. 
Notes: the paratype in the S. S. Berry col- 
lection cannot now be located in the SBMNH, 
and may no longer be extant (P. Valentich- 
Scott to A. К. Kabat, in litt., Mar. 6, 2007). 


pacifica, Chiapaphysa — Taylor, 2003: 170- 


171, fig. 168, pl. 8, fig. 8, map fig. 165. Type 
locality: Rio Tenorito, Hacienda La Pacifica, 
10°29.02’N, 85°9.58’W, Guanacaste, Costa 
Rica. Holotype CAS 114784; paratypes 
MZUCR-INB0003382239. Gastropoda: Pul- 
monata: Physidae. 


Paludiscala Taylor, 1966c: 207. Type species 


Paludiscala caramba Taylor, 1966; original 
designation. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: Taylor questionably 
placed this monotypic genus in the 
Hydrobiidae; Hershler (1985: 58-64) and 
Hershler & Thompson (1992: 85-87) re- 
described this genus and concluded that it 
belonged in the Cochliopinae. 


DWIGHT WILLARD TAYLOR 209 


Paludiscalinae Taylor, 1966c: 207. Described 
as a subfamily in the Hydrobiidae, for Palu- 
discala Taylor, 1966. Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Hershler 
(1984: 65-66) and Hershler & Thompson 
(1992: 86-87) concluded that this subfamily 
was a junior synonym of Cochliopinae 
Stimpson, 1865 (formerly Littoridininae). 
Paupertryonia Taylor, 1987: 38. Type species 
Potamopyrgus cheatumi Pilsbry, 1935; origi- 
nal designation. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: Hershler & Thompson 
(1992: 107) concluded that this genus was 
a junior synonym of Tryonia Stimpson, 1865. 
pecos, Assiminea — Taylor, 1987: 8-9, fig. 2. 
Type locality: seepage at Bitter Lake National 
Wildlife Refuge, 1,250 ft E, 2,100 ft S, sec. 
21, Г. 10$, К. 25E, Chaves Co., New Mexico. 
Holotype LACM 2088; paratypes UTEP 
10051. Gastropoda: Prosobranchia: Assi- 
mineidae. Notes: Taylor (1987) recorded this 
species from three localities in the Pecos 
River basin (near Roswell, New Mexico, and 
near Fort Stockton, Texas), and from the 
Cuatro Ciénegas basin (Coahuila, Mexico). 
Hershler et al. (2007), based on morphologi- 
cal and mitochondrial genetic analyses, de- 
termined that the latter population was not 
conspecific with the former populations, and 
was probably disjunct since the mid-Pleis- 
tocene, if not earlier. Hence, they described 
Assiminea cienegensis as a new species for 
the Cuatro Ciénegas basin population in- 
cluded by Taylor within A. pecos. 
pecosensis, Fontelicella — Taylor, 1987: 27- 
28, fig. 12. Type locality: Blue Spring, center 
SW 1/4 sec. 27, Т. 24$, К. 26E, Eddy Co. 
New Mexico. Holotype LACM 2220; 
paratypes UTEP 10056; ANSP 376021: 
FLMNH 160935, 160933; USNM 854084. 
Gastropoda: Prosobranchia: Hydrobiidae. 
Notes: Hershler (1994: 59) recognized this 
taxon as a valid species of Pyrgulopsis Call 
& Pilsbry, 1886. 

Pecosorbis Taylor, 1985a: 5-6. Type species 
Biomphalaria kansasensis E. G. Berry in E. 
G. Berry & Miller, 1966; original designation. 
Pliocene-Recent. Gastropoda: Pulmonata: 
Planorbidae. 

petenensis, Mayabina — Taylor, 2003: 96-97, 
fig. 77, pl. 3, fig. 10, map fig. 69. Type local- 
ity: Aguado, at NE side of La Libertad, 
16°47.30’N, 90°6.49’W, 200 m, El Petén, 
Guatemala. Holotype CAS 114811; para- 
types CAS 114823; BMNH 20001310. Gas- 
tropoda: Pulmonata: Physidae. 


Phreatomenetus Taylor, 1960b: 60. Described 


as a subgenus of Promenetus F. C. Baker, 
1935. Type species Promenetus umbili- 
catellus (Cockerell, 1887); original designa- 
tion. Gastropoda: Pulmonata: Planorbidae. 


Physellini Taylor, 2003: 167. Described as a 


tribe in the subfamily Physinae, for Physella 
Haldeman, 1843, Chiapaphysa, Costatella, 
Petrophysa, Utahphysa, and Ultraphysella. 
Gastropoda: Pulmonata: Physidae. Notes: 
Taylor (2002a: 25) previously used this name 
as a nomen nudum. 


“Physini” Taylor, 2003: 152. Gastropoda: 


Pulmonata: Physidae. However, this taxon 
was already established as a family-level 
name by Fitzinger (1833), so Taylor is not to 
be credited as the author of this tribe, pur- 
suant to ICZN Article 36.1 (“A name estab- 
lished for a taxon at any rank in the family 
group is deemed to have been simulta- 
neously established for nominal taxa at all 
other ranks in the family group ... The name 
has the same authorship and date at every 
rank.”). 


pinetorum, Fontelicella — Taylor, 1987: 20-21, 


fig. 9. Type locality: spring tributary to Leeds 
Creek, 2,400 ft W, 2,300 ft N, sec. 16, T. 40S, 
R. 14W, Washington Co., Utah. Holotype 
LACM 2217; FLMNH 160946; USNM 
854083. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: Hershler (1994: 44) con- 
cluded that this taxon was a junior synonym 
of Pyrgulopsis kolobensis (Taylor, 1987). 


planispira, Oreoconus — Taylor in McKenna et 


al., 1962: 11-15, figs. 2-4. Type locality: 
USGS Cenozoic loc. 20079: sec. 34, T. 27N, 
R. 97W, Fremont Co., Wyoming. Eocene. 
Holotype USNM 647848. Gastropoda: 
Pulmonata: Bulimulidae. 


Pliopholygidae Taylor, 1966b: 128. New fam- 


ily in the Viviparacea, for Pliopholyx Yen, 
1944. Pliocene. Gastropoda: Prosobranchia: 
Viviparoidea. Notes: Taylor transferred the 
type genus from the Planorbidae 
(Pulmonata) to this new family, and stated 
that this genus had ten species (three de- 
scribed), all from the late Pliocene, Glenns 
Ferry and Cache Valley formations, south- 
ern Idaho and northern Utah. However, the 
seven undescribed species remain manu- 
script names. 


polita, Mayabina — Taylor, 2003: 99-102, figs. 


80-84, pl. 5, figs. 1, 2, map fig. 69. Type lo- 
cality: pasture pool, 50 m W of Rio Tulija, 
1.5 km S of Mexico Highway 186 toward 
Zopo Norte, 17°39.6’N, 92°24.7’W, Tabasco, 


210 KABAT & JOHNSON 


Mexico. Holotype CAS 114783; paratypes 
CAS 114817; BMNH 20001311; ZIBM 
CMNO 1160. Gastropoda: Pulmonata: Phy- 
sidae. 

polyskelidis, Lutrilimnea — Taylor in Taylor & 
Smith, 1981: 361-363, pls. 5-7. Type local- 
ity: limestone and coquina beds, SW 1/4 sec. 
17, Г. 23N, R. 29E, strike N 60° W, dip 4° NE, 
thickness 15-20 ft, weathering on the service 
1100-1300 ft E, 1300-1400 ft N of the SW 
section corner, Mopung Hills, Churchill Co., 
Nevada. Pliocene. Holotype UMMZ 250106. 
Gastropoda: Pulmonata: Lymnaeidae. 

Potamopyrgue Taylor, 1987: 38. Error for 
Potamopyrgus Stimpson, 1865. 

quadripaludium, Mexithauma — Taylor, 1966c: 
205-207, pl. 19, figs. 58-63, text fig. 22. Type 
locality: Laguna Tio Candido, 14 km $ of 
Cuatro Ciénegas, Coahuila, Mexico. Holo- 
type UMMZ 220214. Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Hershler 
(1985: 72-78) redescribed this taxon. 

rexroadensis, Polygyra (Erymodon) — Taylor, 
1960b 462 ple 1 Mos. 14 23, pl2 0525: 
Type locality: Fox Canyon, Univ. Michigan 
loc. UM-K1-47, Sec. 35, T. 34$, К. 30W, a 
lenticular bed of blue-gray clay, 10 ft below 
the caliche, Meade Co., Kansas. Late 
Pliocene. Holotype UMMZ 177610-a; 
paratypes UMMZ 177610, 183050, 183082. 
Gastropoda: Pulmonata: Polygyridae. 

roswellensis, Fontelicella — Taylor, 1987: 14— 
16, fig. 6. Type locality: seepage 1,250 ft E, 
2,100 ft S, sec. 21, T. 10S, R. 25E, Chaves 
Co., New Mexico. Holotype LACM 2213; 
paratypes UTEP 10057. Gastropoda: Proso- 
branchia: Hydrobiidae. Notes: Hershler 
(1994: 63) recognized this taxon as a valid 
species of Pyrgulopsis Call & Pilsbry, 1886. 

sanctijohannis, Mayabina — Taylor, 2003: 102— 
104, fig. 85, pl. 3, figs. 5, 6, map figs. 15, 69. 
Type locality: Barra del Colorado, 
10°46.37’N, 83°35.27’W, Limón, Costa Rica. 
Holotype CAS 114790; paratypes CAS 
114780; BMNH 20001312; MZUCR- 
INBO003382237; MZUCR 69-1. Gastropoda: 
Pulmonata: Physidae. 

sanguinichristi, Pisidium (Cyclocalyx) — Tay- 
lor, 1987: 47-48, fig. 24. Type locality: Middle 
Fork Lake, a cirque lake at 10,845 ft (3,306 
m) elevation, Sangre de Cristo Mountains, 
Taos Co., New Mexico. Holotype LACM 
2258. Bivalvia: Sphaeriidae. 

Savaginius Taylor, 1966b: 130. Type species 
Paludestrina nanna Chamberlin & Berry, 
1933; original designation. Pliocene and 
Pleistocene. Gastropoda: Prosobranchia: 


Hydrobiidae. Notes: Hershler (1994: 5, 13) 
concluded that this taxon was a junior syn- 
onym of Pyrgulopsis Call & Pilsbry, 1886. 

scaevoscala, Gastrocopta (Gastrocopta) — 
Taylor, 1960b: 70, pl. 1, figs. 33, 34. Type 
locality: along the banks of a tributary of 
Stump Arroyo, SE 1/4 SW 1/4 and SW 1/4 
SE 1/4 sec. 22, Т. 335, К. 29W, Meade Co., 
Kansas. Bender local fauna, Late Pliocene 
and early Pleistocene. Holotype UMMZ 
184320; paratypes UMMZ 184125, 183017. 
Gastropoda: Pulmonata: Pupillidae. 

shotwelli, Carinifex — Taylor, 1963a: 40, figs. 
19-36. USGS loc. 21173, Malheur Co., Or- 
egon. Black Butte local fauna, Juntura For- 
mation, Miocene. Holotype USNM 563110; 
paratypes USNM 563111, 563112, 563113, 
563114. Gastropoda: Pulmonata: Planor- 
bidae. 

sinaloae, Ultraphysella — Taylor, 2003: 192, 
figs. 187—191, pl. 9, fig. 1, map fig. 15. Type 
locality: pool at road 2.5 mi from Villa Union 
toward Siquerios, 23°13.4’N, 106°12.5 W, 
Sinaloa, Mexico. Holotype CAS 146096. 
Gastropoda: Pulmonata: Physidae. 

sinusdulcensis, Tropinauta — Taylor, 2003: 111, 
figs. 91-94. Type locality: small stream in 
pasture 3 km SE of Golfito, 8°36.68'N, 
83°8.48’W, Puntarenas, Costa Rica. Holo- 
type CAS 146095; paratypes MZUCR- 
INBO003382246. Gastropoda: Pulmonata: 
Physidae. 

skinneri, Physa — Taylor, 1954d: 9. Type lo- 
cality: SE corner of sec. 6, T. 5N, R. 28E, 
Beaver Co., Oklahoma. Pleistocene, prob- 
ably Illinoian age, Berends fauna. Holotype 
UMMZ 181292; paratypes UMMZ 177533; 
MCZ 198177; USNM 562010. Gastropoda: 
Pulmonata: Physidae. Notes: Taylor (1988: 
45-55, fig. 1; 2003: 166-167, fig. 156) re- 
described this species. 

sonomae, Archiphysa — Taylor, 2003: 183- 
184, pl. 10, figs. 4, 7, map fig. 176. Type lo- 
cality: artificial pond, 2,500 ft S, 4,300 ft W, 
sec. 30, T. 9N, R. 9W, Sonoma Co., Califor- 
nia. Holotype CAS 114807; paratypes CAS 
114803. Gastropoda: Pulmonata: Physidae. 

spathidophallus, Stenophysa — Taylor, 2003: 
121-123, figs. 110-116, pl. 8, fig. 9. Type 
locality: ditch from Seletar Reservoir, 100 m 
W of Upper Thompson Road, Singapore. 
Holotype CAS 114804. Gastropoda: Pulmo- 
nata: Physidae. Notes: Taylor (2003: 123) 
concluded that this species was probably 
“transported through the trade in tropical 
fish,” and that “| speculate it may [actually] 
occur in northeastern South America.” 


DWIGHT WILLARD TAYLOR 201 


Stenophysini Taylor, 2003: 111. New tribe in 
the subfamily Aplexinae, for Stenophysa 
Martens, 1898, Afrophysa Starobogatov, 
1967, and a “name uncertain” group in Ar- 
gentina and perhaps adjacent countries. 
Gastropoda: Pulmonata: Physidae. Notes: 
Taylor (2002a: 25) previously used this name 
as a nomen nudum. 

stocktonensis, Tryonia — Taylor, 1987: 37-38, 
fig. 18. Type locality: Diamond Y Draw, 9 mi 
N of Fort Stockton and 0.5 mi W of State 
Highway 18, Pecos Co., Texas. Holotype 
LACM 2090; paratypes UTEP 10065; ANSP 
376030; FLMNH 160947; USNM 854092. 
Gastropoda: Prosobranchia: Physidae. 
Notes: Hershler & Thompson (1992: 110) 
concluded that this taxon was a junior syn- 
onym of Tryonia circumstriata Leonard & Ho, 
1960. 

tempisquensis, Mayabina — Taylor, 2003: 109, 
fig. 90, pl. 4, fig. 4, map figs. 15, 68. Type 
locality: Parque Nacional Palo Verde, edge 
of marshes 100 m E of W end of airstrip, 
10°20.68’N, 85°20.60’W, Guanacaste, Costa 
Rica. Holotype CAS 146092; paratypes 
MZUCR 70-01, MZUCR-INB0003382244. 
Gastropoda: Pulmonata: Physidae. 

thermalis, Fontelicella — Taylor, 1987: 28-30, 
fig. 13. Type locality: hot spring on E side of 
Gila River, NE 1/4 SW 1/4 sec. 17, Т. 13S, 
R. 13W, Grant Co., New Mexico. Holotype 
LACM 2224: paratypes UTEP 10058; ANSP 
376026; FLMNH 160941; USNM 854086. 
Gastropoda: Prosobranchia: Hydrobiidae. 
Notes: Hershler (1994: 66) recognized this 
taxon as a valid species of Pyrgulopsis Call 
& Pilsbry, 1886. 

trivialis, Fontelicella — Taylor, 1987: 30-32, fig. 
14. Type locality: spring-fed pond, 1,000 ft 
N of SW corner, sec. 5, T. 5N, R. 29E, Apache 
Co., Arizona. Holotype LACM 2225; para- 
types UTEP 10059; ANSP 376022; FLMNH 
160945; USNM 854080. Gastropoda: 
Prosobranchia: Hydrobiidae. Notes: 
Pyrgulopsis confluentis Hershler in Hershler 
& Landye, 1988: 32-35 is a junior subjective 
synonym of Fontelicella trivialis Taylor, 1987 
(fide Hershler & Landye, 1988: 58). Hershler 
(1994: 68) recognized this taxon as a valid 
species of Pyrgulopsis Call & Pilsbry, 1886. 

Tropinauta Taylor, 2003: 110-111. Type spe- 
cies Tropinauta sinusdulcensis Taylor, 2003; 
Original designation. Gastropoda: 
Pulmonata: Physidae. Notes: Taylor (2002a: 
25) previously used this name as a nomen 
nudum. 


Ultraphysella Taylor, 2003: 191. Type species 
Ultraphysella sinaloae Taylor, 2003; original 
designation. Gastropoda: Pulmonata: 
Physidae. Notes: Taylor (2002a: 25) previ- 
ously used this name as a nomen nudum. 

ursina, Lutrilimnea — Taylor in Taylor & Smith, 
1981: 364-366, pl. 9. Type locality: drift on 
shore at S end of Bear Lake, SW 1/4 sec. 
23, T. 13N, R. 5E, Rich Co., Utah. Holocene 
(“became extinct about 8,000 years ago”). 
Holotype UMMZ 250113. Gastropoda: 
Pulmonata: Lymnaeidae. 

Utahphysa Taylor, 2003: 175-177. Type species 
Aplexa microstriata Chamberlin & Berry, 1930; 
original designation. Gastropoda: Pulmonata: 
Physidae. Notes: Taylor (2002a: 25) previously 
used this name as a nomen nudum. 

vernalis, Physa — Taylor & Jokinen, 1984: 190, 
figs. 1-11. Type locality: Bluebird Pond, site 
no. 240, Windham, Windham Co., Connecti- 
cut. Holotype MCZ 294071. Gastropoda: 
Pulmonata: Physidae. Notes: Taylor (2003: 
157, figs. 153-155, pl. 8, fig. 2, map fig. 147) 
transferred this species to Laurentiphysa 
Taylor, 2003. 

wilsoni, Planorbella (Seminolina) — Taylor, 
1966b: 111-113, pl. 8, figs. 7-9. Type local- 
ity: USGS Cenozoic loc. 22704, “unitA,” Belle 
Glade, Palm Beach Co., Florida. Late Plio- 
cene or early Pleistocene. Holotype USNM 
644835. Gastropoda: Pulmonata: Planorbi- 
dae. 

Yaquicoccus Taylor, 1987: 34. Type species 
Yaquicoccus bernardinus Taylor, 1987; origi- 
nal designation. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: Hershler (1994: 5, 13) 
concluded that this taxon was a junior syn- 
onym of Pyrgulopsis Call & Pilsbry, 1886. 


Eponyms 


Taylorconcha Hershler et al., 1994: 233. Type 
species Taylorconcha serpenticola Hershler 
et al., 1994. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: “Honoring Dwight Tay- 
lor, for his discovery and early work on this 
genus and, more generally, in recognition of 
his lifetime of fieldwork and research on the 
systematics, biology, and biogeography of 
the freshwater molluscan fauna of western 
North America.” The type species, commonly 
known as the “Bliss Rapids Snail,” was rec- 
ommended by Taylor in his 1982 report to 
the USFWS for listing as an endangered 
species, and was subsequently listed as a 
threatened species. 


2412 KABAT & JOHNSON 


taylori, Hadoceras — Hershler & Longley, 1986: 
121-136. Gastropoda: Prosobranchia: 
Hydrobiidae. Notes: “Named after Dwight 
Taylor in recognition for his discovery of this 
species and immense contributions to the 
study of west American freshwater mollusks.” 

taylori, Helminthoglypta (Coyote) — Reeder & 
Roth, 1988: 254-256. Gastropoda: Pulmo- 
nata: Helminthoglyptidae. 

taylori, Pyrgulopsis — Hershler, 1995: 363-364. 
Gastropoda: Prosobranchia: Hydrobiidae. 
Notes: “This species is named in honor of 
Dwight W. Taylor, in recognition of his many 
years of fieldwork and associated research 
on hydrobiids both in California and through- 
out the western United States.” 

taylori, Radiocentrum — Roth, 1986: 249. Gas- 
tropoda: Pulmonata: Punctoidea. 


Bibliography of Taylor’s Publications 


The publications are listed in chronological 
order, without regard to senior authorship of 
multiple-authored papers. An asterisk (*) indi- 
cates publications with no new taxa. Unpub- 
lished agency reports and other manuscripts 
are not included herein. 

The University of California Berkeley Library 
has a two-volume bound set of Taylor’s pa- 
pers from 1950 to 1966, evidently bound from 
the reprints that he donated to the Museum of 
Paleontology. However, these volumes do not 
include all of his publications from that period. 


* TAYLOR, D. W., 1949a [19 March], STS win- 
ner writes. Science News Letter 55(12): 182 
[excerpt of Taylor 1949b]. 

* TAYLOR, D. W., 1949b [November], À mala- 
cological survey of Nantucket Island, Mas- 
sachusetts. Pp. 18-21 and back cover, in: 
How you can search for science talent: a 
book of facts about the Ninth Annual Science 
Talent Search for Westinghouse Science 
Scholarships. Washington, D.C., Science 
Clubs of America/Science Service. 

TAYLOR, D. W., 1950 [11 January], Three new 
Pyrgulopsis from the Colorado Desert, Cali- 
fornia. Leaflets in Malacology, 1(7): 27-33. 

* TAYLOR, D. W., 1952 [10 July], Notes on 
the freshwater mollusks of Yellowstone Park, 
Wyoming. Leaflets т Malacology, 1(9): 43- 
49, 1 pl. 

TAYLOR, D. W., 1954a [10 January], À new 
Promenetus (Planorbidae) from Panama. 
Revista de la Sociedad Malacologica “Carlos 
de la Torre” [Havana], 9(2): 37-38. 


* TAYLOR, D. W., 1954b [May], Some Late 
Cenozoic Molluscan Faunas from Kansas 
and Nebraska. Master’s Thesis, University 
of California, Berkeley. 188 pp. [unpub- 
lished]. 

TAYLOR, D. W., 1954c [June], Nonmarine 
molluscs from Barstow Formation of south- 
ern California. United States Geological Sur- 
vey, Professional Paper, 254C: 67-80, plate 
20: 

TAYLOR, D. W., 19544 [13 August], A new 
Pleistocene fauna and new species of fossil 
snails from the High Plains. Occasional Pa- 
pers, Museum of Zoology, University of 
Michigan, 577: 1-16. 

* TAYLOR, D. W. & С. W. HIBBARD, 1955, A 
new Pleistocene fauna from Harper County, 
Oklahoma. Oklahoma Geological Survey, 
Circular, 37: 1-23. 

* TAYLOR, D. W., 1956, Pliocene mollusks 
from Jackson Hole, Grand Valley, and Star 
Valley, Wyoming and Idaho. Pp. 123-125, 1 
pl., in: Guidebook, Wyoming Geological As- 
sociation, 11th Annual Field Conference, 
1956, Jackson Hole. 256 pp. 

* TAYLOR, D. W., 1957a [January], Late Ceno- 
zoic paleoecology and molluscan faunas of 
the High Plains. Ph.D. Thesis, University of 
California, Berkeley. vii + 351 pp. [unpub- 
lished; not seen; listed by title only in “Index 
to American Doctoral Dissertations 1956- 
1957 р: 1021958). 

TAYLOR, D. W., 1957b [13 June], Pliocene 
fresh-water mollusks from Navajo County, 
Arizona. Journal of Paleontology, 31(3): 654— 
661. 

* ROBINSON, С. D., С. Е. LEWIS & D. W. TAY- 
LOR, 1957 [December], Eocene continental 
deposits in Three Forks Basin, Montana. 
Geological Society of America Bulletin, 
68(12) [part 2]: 1786 [abstract of paper pre- 
sented at GSA meeting, Atlantic City, New 
Jersey, 1-3 November 1957]. 

* HERRINGTON, Н. В. & D. W. TAYLOR, 1958 
[15 August], Pliocene and Pleistocene 
Sphaeriidae (Pelecypoda) from the central 
United States. Occasional Papers, Museum 
of Zoology, University of Michigan, 596: 1— 
128, 1 pl. [new taxa by Herrington alone]. 

* TAYLOR, D. W., 1958 [12 December], Geo- 
logic range and relationships of the fresh- 
water snail Anisus pattersoni. Journal of 
Paleontology, 32(6): 1149-1153. 

* TAYLOR, D. W., 1960a [April?], Distribution 
of the freshwater clam Pisidium ultra- 
montanum; a zoogeographic inquiry. Ameri- 


DWIGHT WILLARD TAYLOR 213 


can Journal of Science, 258A: 325-334, 1 
pl. + errata [note: MCZ issue received on 25 
April 1960; USNM issue received on 27 April 
1960]. 

* HIBBARD, C. W. & D. W. TAYLOR, 1960 [1 
July], Two Late Pleistocene faunas from 
southwestern Kansas. Contributions from 
the Museum of Paleontology, The University 
of Michigan, 16(1): 1-223, pls. 1-16 [new 
mammalian taxa by Hibbard alone]. 

TAYLOR, D. W., 1960b [July?], Late Cenozoic 
molluscan faunas from the High Plains. 
United States Geological Survey Profes- 
sional Paper, 337: iv + 94 pp., pls. 1—4 [note: 
Library of Congress issue received on 7 July 
1960; Univ. of Washington issue received on 
21 July 1960]. 

* TAYLOR, D. W., 1961a [1 April], The fresh- 
water clam Pisidium ultramontanum Prime 
in Modoc County, California. The Veliger, 
3(4): 111. 

* TAYLOR, D. W., 1961b [17 November], Com- 
ments on the proposed suppression of Palu- 
dina lustrica Say, 1821. Z.N.(S.) 730. Bulletin 
of Zoological Nomenclature, 18(6): 379. 

* TAYLOR, D. W. & Н. В. HERRINGTON, 1962 
[1 January], The freshwater clam Pisidium 
tremperi (Hannibal). The Veliger, 4(3): 129— 
131, plate 28. 

McKENNA, M. C., P. ROBINSON & D. W. 
TAYLOR, 1962 [12 September], Notes on 
Eocene Mammalia and Mollusca from Tab- 
ernacle Butte, Wyoming. American Museum 
Novitates, 2102: 1-33 [description of mol- 
lusks by Taylor alone]. 

* LOVE, J. D. & D. W. TAYLOR, 1962 [No- 
vember?], Faulted Pleistocene strata near 
Jackson, northwestern Wyoming. United 
States Geological Survey, Professional Pa- 
per, 450-D: 2136-0139 [note: Library of 
Congress issue received on 8 November 
1962; Univ. of Washington issue received on 
26 November 1962]. 

* TAYLOR, D. W. & N. F. ЗОНЕ, 1962 [14 No- 
vember], An outline of gastropod classifica- 
tion. Malacologia, 1(1): 7-32. 

TAYLOR, D. W., 1963a [April], Mollusks of the 
Black Butte local fauna. Pages 35—41, in: J. 
A. SHOTWELL, ed., The Juntura Basin: stud- 
ies in earth history and paleoecology. Trans- 
actions of the American Philosophical 
Society, (n.s.) 53(1): 1-77. 

* TAYLOR; DW, AIS WALTER. JB, 
BURCH, 1963 [7 August], Freshwater snails 
of the subgenus Hinkleyia (Lymnaeidae: 
Stagnicola) from the western United States. 
Malacologia, 1(2): 237-281, 4 pls. 


* RUBIN, M. & D. W. TAYLOR, 1963 [16 Au- 
gust], Radiocarbon activity of shells from liv- 
ing clams and snails. Science, 141(3581): 
687. 

* TAYLOR, D. W., 1963b [1 October], Errone- 
ous records of freshwater clams 
(Spaheriidae) from California. The Veliger, 
GTR 

* TAYLOR, D. W., 1964 [November], Histori- 
cal analysis of distribution of west American 
freshwater molluscs. American Zoologist, 
4(4): 436-437 [Abstract of paper presented 
at American Society of Zoologists meeting, 
Knoxville, Tennessee, 28 December 1964]. 

* McCULLOCH, D. $., D. W. TAYLOR & M. 
RUBIN, 1965 [May], Stratigraphy, non-ma- 
rine mollusks, and radiometric dates from 
Quaternary deposits in the Kotzebue Sound 
area, western Alaska. Journal of Geology, 
73(3): 442—453. 

GREGG WW. Où& В. We TAYLOR. 1905191 
August], Fontelicella (Prosobranchia: 
Hydrobiidae), a new genus of west Ameri- 
can freshwater snails. Malacologia, 3(1): 
103-110. 

* TAYLOR, D. W., 1965, The study of Pleis- 
tocene nonmarine mollusks in North 
America. Pp. 597-611, т: Н.Е. WRIGHT & D. 
G. FREY, eds., The Quaternary of the United 
States; a Review Volume for the VII Con- 
gress of the International Association for 
Quaternary Research. Princeton, New Jer- 
sey, Princeton University Press. x + 922 pp. 

* HIBBARD, C. W., C. E. RAY, D. E. SAVAGE, 
О. W. TAYLOR 4 J. E. GUILDAY, 1965, Qua- 
ternary mammals of North America. Pp. 509- 
525, in: H.E. WRIGHT & D. G. FREY, eds., The 
Quaternary of the United States; a Review 
Volume for the VII Congress of the Interna- 
tional Association for Quaternary Research. 
Princeton, New Jersey, Princeton University 
Press. x + 922 pp. 

* TAYLOR, D. W., 1966a [1 January], An east- 
ern American freshwater mussel, Anodonta, 
introduced into Arizona. The Veliger, 8(3): 
197—198, plate 28. 

* TAYLOR, D. W. & T. UYENO, 1966 [20 Janu- 
ary] ['December 1965”], Evolution of host 
specificity of freshwater salmonid fishes and 
mussels in the North Pacific region. Venus, 
24(3): 199-209 [in Japanese; English ab- 
stract]. 

TAYLOR, D. W., 1966b [18 August], Summary 
of North American Blancan nonmarine mol- 
lusks. Malacologia, 4(1): 1-172, pls. 1-8. 
[Note: this journal issue contained three “pa- 
pers planned for the VII Congress of the In- 


214 KABAT & JOHNSON 


ternational Association for Quaternary Re- 
search”). 

* TAYLOR, D. W. & W. L. MINCKLEY, 1966 
[August] [“September—October’], New world 
for biologists. Pacific Discovery, 19(5): 18— 
22, 6 pls., 1 map [Library of Congress issue 
received 29 August 1966; USNM issue re- 
ceived on 8 Sept. 1966]. 

TAYLOR, D. W., 1966c [1 October], Aremark- 
able snail fauna from Coahuila, Mexico. The 
Veliger, 9(2): 152-228, pls. 8-19. 

* TAYLOR, D. W., 1966d [1 October], Review 
ГА preliminary checklist of invertebrates col- 
lected from Lake Tahoe, 1961-1964, by Ted 
C. Frantz and Almo J. Cardone. Biological 
Society of Nevada, Occasional Papers, 8: 1— 
12 (15 January, 1966)']. The Veliger, 9(2): 253. 

* TAYLOR, D. W., 1967a [22 February] ['De- 
cember 1, 1966”], A remarkable snail fauna 
from Coahuila, Mexico. Report of the Ameri- 
can Malacological Union for 1966: 70-72 
[Abstract of paper presented at meeting of 
the AMU-Pacific Division, Seattle, Washing- 
ton, 19-22 June 1966]. 

* TAYLOR, D. W., 1967b [4 August], Fresh- 
water clam Sphaerium transversum (Say) in 
Arizona. Southwestern Naturalist, 12(2): 
202-203. 

* TAYLOR, D. W., 1967c [1 October], Fresh- 
water mollusks collected by the United States 
and Mexican Boundary Surveys. The Veliger, 
10(2): 152-158. 

* TAYLOR, D. W., 1967d [October], Late Pleis- 
tocene molluscan shells from the Tule 
Springs area. In: H. M. WORMINGTON & D. 
ELLIS, eds., Pleistocene studies in southern 
Nevada. Nevada State Museum Anthropo- 
logical Papers, 13: 395-399. 

* TAYLOR, D. W., 1968 [20 March], Late Pleis- 
tocene nonmarine mollusks from the State 
of Puebla, Mexico. Annual Report of the 
American Malacological Union, 34: 76-78 
[Abstract of paper that “was announced but 
was not presented” at meeting of the AMU- 
Pacific Division, Pacific Grove, California, 28 
June — 1 July 1967 (Hanselman, 1968: 66)]. 

* LONGeG.E: 4D, Wr TAYLOR, 1970 [9 
March], Estuarine mollusks of the Cholla Bay, 
Sonora, Mexico. The Echo (Western Soci- 
ety of Malacologists), 2: 17-18, 39 [Abstract 
of paper presented at meeting of the WSM, 
Pacific Grove, California, 18-21 June 1969]. 

* TAYLOR, D. W., 1970a [30 June], West 
American freshwater Mollusca, 1: bibliogra- 
phy of Pleistocene and Recent species. 
Memoirs, San Diego Society of Natural His- 
tory, 4: 1-73, 1 plate. 


* TAYLOR, D. W., 1970b [14 November], Sym- 
posium on the rare and endangered mollusks 
of North America, 4. Western freshwater mol- 
lusks. Malacologia, 10(1): 33-34 [With sum- 
mary of Taylor's manuscript by Harold D. 
Murray; Taylor did not attend this symposium 
(The American Malacological Union, Inc., 
Annual Reports for 1968, Bulletin, 35: 3)]. 

* TAYLOR, D. W. &A. G. SMITH, 1971 [1 April], 
Harold Hannibal (1889-1965) with a review 
of his molluscan research. The Veliger, 13(4): 
303-316, 5 pls. 

* TAYLOR, D. W., 1974 [15 July], The Tertiary 
gastropod Orygoceras found living. Archiv 
fur Molluskenkunde, 104(1-3): 93-96. [Re- 
printed (1974), with Hungarian title, “Eld 
harmadkori Orygocerasok,” and Hungarian 
abstract, Soosiana, 2: 37—44]. 

* TAYLOR, D. W., 1975, Index and bibliogra- 
phy of Late Cenozoic freshwater Mollusca 
of western North America. Claude W. 
Hibbard Memorial Volume 1. Papers on Pa- 
leontology (Museum of Paleontology, Univer- 
sity of Michigan), 10: 1-384 [one-page errata 
issued in 1976]. 

* TAYLOR, D. W., 1978 [February], Comments 
on the proposed designation of a type-spe- 
cies for Pleurocera Rafinesque, 1818. 
Z.N.(S.) 83. Bulletin of Zoological Nomen- 
clature, 34(4): 199. 

* TAYLOR, D. W., 1981 [July], Freshwater mol- 
lusks of California: a distributional checklist. 
California Fish and Game, 67(3): 140-163. 

TAYLOR, D. W. & G. R. SMITH, 1981 [31 De- 
cember], Pliocene molluscs and fishes from 
northeastern California and northwestern 
Nevada. Contributions from the Museum of 
Paleontology, The University of Michigan, 
25(18): 339-413, 19 pls. 

TAYLOR, D. W., 1983 [81 December], Late 
Tertiary mollusks from the lower Colorado 
River Valley. Contributions from the Museum 
of Paleontology, The University of Michigan, 
26(13): 289-298. 

TAYLOR, D. W. & Е. H. JOKINEN, 1984 [5 
November], A new species of freshwater 
snail (Physa) from seasonal habitats in Con- 
necticut. Freshwater Invertebrate Biology, 
3(4): 189-202. 

TAYLOR, D. W., 1985a [February], Pecosorbis, 
a new genus of fresh-water snails (Planor- 
bidae) from New Mexico. New Mexico Bu- 
reau of Mines & Mineral Resources, Circular, 
194: 1-17. 

* TAYLOR, D. W., 1985b, Miocene freshwater 
mollusks from the Clarkia fossil site, Idaho. 
Pp. 73-74, in: С. J. SMILEY, ed., Late Ceno- 


DWIGHT WILLARD TAYLOR 215 


zoic history of the Pacific Northwest: Inter- 
disciplinary Studies on the Clarkia Fossil 
Beds of Northern Idaho. American Associa- 
tion for the Advancement of Science, Pacific 
Division, San Francisco. 417 pp. 

* TAYLOR, D. W., 1985c, Evolution of freshwa- 
ter drainages and molluscs in western North 
America. Pp. 265-321, in: С. J. SMILEY, ed., 
Late Cenozoic history of the Pacific North- 
west: Interdisciplinary Studies on the Clarkia 
Fossil Beds of Northern Idaho. American As- 
sociation for the Advancement of Science, 
Pacific Division, San Francisco. 417 pp. 

* CONEY ©. С. & DW. TAYLOR; 1986 [31 
January] [*1985”], Systematic position of 
Quincuncina mitchelli (Simpson, 1896). An- 
nual Report, Western Society of Malacolo- 
gists, 18: 12-13 [Abstract revised November 
12, 1985, based on new evidence”] [Abstract 
of paper presented by Coney at WSM meet- 
ing, Santa Barbara, California, 18—21 August 
1985, which Taylor did not attend; after the 
meeting, Coney added Taylor as a co-author]. 

* TAYLOR, D. W., 1986, Fossil molluscs from 
the Lake Hill archaeological site, Panamint 
Valley, southeastern California. Contributions 
of the Great Basin Foundation (San Diego), 
2: 42-54. 

TAYLOR, D. W., 1987 [September], Fresh- 
water molluscs from New Mexico and vicin- 
ity. New Mexico Bureau of Mines & Mineral 
Resources, Bulletin, 116: iv + 5—50. 

* TAYLOR, D. We & В, С. BRIGHT,--1987, 
Drainage history of the Bonneville Basin. 
Utah Geological Association, Publication, 16: 
239-256 [paper presented at symposium, 
“Cenozoic geology of western Utah”, Lake 
City, Utah, 23-26 September 1987]. 

* TAYLOR, D. W., 1988a [January], Phylum: 
Mollusca. Pp. 32-57, in: J. GRAY, ed., Evolu- 
tion of the freshwater ecosystem: the fossil 
record. Palaeogeography, Palaeoclimatology, 
Palaeoecology, 62(14): 1-214. 

* TAYLOR, D. W. & J. GRAY, 1988 [January], 
Class: Mammalia. Pp. 165-175, in: J. GRAY, 
ed., Evolution of the freshwater ecosystem: 
the fossil record. Palaeogeography, Palaeo- 
climatology, Palaeoecology, 62(1-4): 1-214. 

* TAYLOR, D. W., 1988b [January], Aspects 
of freshwater mollusc ecological biogeogra- 
phy. Palaeogeography, Palaeoclimatology, 
Palaeoecology, 62(1-4): 511-576. 

TAYLOR, D. W., 1988c [2 December], New 
species of Physa (Gastropoda: Hygrophila) 
from the western United States. Malacologi- 
cal Review, 21(1-2): 43-79. 

* BOUCOT, A. J., H. A. McCLUER, Е. ALVA- 
REZ, JR, P ROSS, DEM TAYLOR, W. 


STRUVE, N. N. SAVAGE & S. TURNER, 
1989 [25 September], New Devonian fossils 
from Saudi Arabia and their biogeographic 
affinities. Senckenbergiana Lethaea, 69(5- 
6): 535-597 [Taylor, “Nonmarine molluscs,” 
рр. 557—559]. 

* GRAY, J. & D. W. TAYLOR, 1992 [May], Late 

Tertiary environmental change: pollen and 

mollusc evidence from the Pacific Northwest. 

Geological Society of America, Abstracts 

with Programs, 24(5): 28 [abstract of paper 

presented at the meeting of the Cordilleran 

Section, GSA, Eugene, Oregon, 11-13 May 

1992]. 

TAYLOR, D. W., 1994 [June] [“December 

1993”], Moluscos dulceacuicolas de Costa 

Rico: introduccion y lista preliminar. Revista 

de Biología Tropical (San José, Costa Rica), 

41(3A): 653-655 [in Spanish; English ab- 
stract]. 

* TAYLOR, D. W., 1997a [9 January], An old 
new species of Polymesoda (Bivalvia, Corbi- 
culidae) from the Pacific coast of Mexico. 
The Festivus, 29(1): 3-5. 

TAYLOR, D. W., 1997b [1 August], A new 
mussel, Disconaias conchos (Bivalvia: 
Unionidae) from Rio Conchos of the Rio 
Grande System, Mexico. Occasional Papers 
on Mollusks, 5(75): 419-425. 

* TAYLOR, D. W., 2002a, New data on bioge- 
ography, classification and phylogeny of Phy- 
sidae (Gastropoda: Hygrophila). Visnyk 
Zhytomyrskoho Pedahohichnoho Univer- 
sytetu [Proceedings of the lvan Franko Zhyto- 
myr Pedagogical University] (Zhytomyr), 10: 
24—26 [proceedings of malacological confer- 
ence held in Zhytomyr, Ukraine, 13-15 Мау 
2002]. 

* TAYLOR, D. W., 2002b [*1998-1999”], 
Harold William Harry, 1921-1995. Malaco- 
logical Review, 31/32: 159-163. 

TAYLOR, D. W., 2003 [March], Introduction to 
Physidae (Gastropoda: Hygrophila); bioge- 
ography, classification, morphology. Revista 
de Biología Tropical, International Journal of 
Tropical Biology and Conservation (San 
José, Costa Rica), 51 (Supplement 1): viii + 
287 DD. 

* TAYLOR, D. W., 2004a [August], Morpho- 
logical revision of freshwater snails, Family 
Physidae / Revision morfolögica de caraco- 
les dulciacuicolas, Familia Physidae. 
Comunicaciones de la Socieded Malacolo- 
gica del Uruguay, 8(82-83): 279-282. 

* TAYLOR, D. W., 2004b [October], Trans-Pa- 
cific relationships in Physidae (Gastropoda, 
Pulmonata). Pp. 155-156, in: O. YA. SEMENI- 
KHINA, ed., Abstracts of the conference “Mol- 


+ 


216 KABAT & JOHNSON 


lusks of the Northeastern Asia and Northern 
Pacific: Biodiversity, Ecology, Biogeography, 
and Fauna History,” October 4-6, 2004, 
Vladivostok, Russia. Vladivostok, Dalnauka. 
176 pp. [abstract] [Taylor did not attend this 
meeting (K. A. Lutaenko to A. R. Kabat, in 
litt., May 21, 2007)]. 


SECONDARY LITERATURY 


ADLER, K. K., 2007 [11 July], Herpetologists of 
the past, Part 2. Pp. 7-273, in: K. K. ADLER, ed., 
Contributions to the history of herpetology, Vol. 
2. St. Louis, Missouri, Society for the Study of 
Amphibians and Reptiles. 389 pp. 

ANONYMOUS, 1948 [11 December], Young 
scientists compete. Science News Letter 
55(11): 380. 

ANONYMOUS, 1949a [26 February], California 
youth studies New England island life. Science 
News Letter, 55(9): 134-135. 

ANONYMOUS, 1949b [11 March], [News]. Sci- 
ence, 109(2828): 270. 

ANONYMOUS, 1949c [12 March], Science Tal- 
ent Institute: President Truman promised the 
40 honor-trip winners that when peace came 
to the world there would be an immense num- 
bers of jobs in the field of science. Science 
News Letter, 55(11): cover, 165. 

ANONYMOUS, 1949d [19 March], Top science 
scholarships: surveyor of shellfish life off east- 
ern coast, Dwight W. Taylor, received $2,800 
award. Science News Letter, 55(12): cover, 
178-181. 

ANONYMOUS, 1949e [21 March], Top of the 
crop. Time, 53(12): 49. 

ANONYMOUS, 1955, Grace Wyatt 1894-1954. 
The Nantucket Maria Mitchell Association, An- 
nual Report, 53: 12. 

ANONYMOUS, 1957 [19 October], Geology: 
examine snail shells for past climate changes. 
Science News Letter, 72(16): 249. 

ANONYMOUS, 1966a, Mrs. Willard, 92 dies; 
family owned hotel. The Evening Star (Wash- 
ington, D.C.), March 16, p. B-6. 

ANONYMOUS, 1966b, Helen Willard dies here 
at 92. Washington Post, March 17, p. B-6. 

ANONYMOUS, 2007 [March], Dwight Willard 
Taylor ‘49, January 18, 1932 — August 3, 2006, 
Memorial Service. Claremont (California), 
Webb School. 8 pp. 

BARRIENTOS, Z. & M. SPRINGER, 2007 [March], 
In memoriam: Dwight Willard Taylor, “Don 
Guillermo”. Revista de Biologia Tropical, Inter- 
national Journal of Tropical Biology and Con- 
servation (San José, Costa Rica), 55(1): ix—xii. 

BELL, S. K., 2004, Malcolm C. McKenna: a bi- 
ography and bibliography. Bulletin of the Ameri- 
can Museum of Natural History, 285: 11-24. 

BURCH, J. B. & J. M. HUBER, 1968 [20 March], 
Malacologia — five years of publication. The 
American Malacological Union, Inc., Annual 
Reports for 1967: 29-30. 

CLARK, B. L., 1915 [30 August], Fauna of the 
San Pablo Group of middle California. Univer- 


sity of California Publications, Bulletin of the 
Department of Geology, 8(22): 385-572, pls. 
42-71. 

COLE, G. A., 1984, Crustacea from the Bolsön 
of Cuatro Ciénegas, Coahuila, Mexico. Jour- 
nal of the Arizona-Nevada Academy of Sci- 
ence, 19(1): 3-12. 

COLLINS, J. P., J. DEACON, T. DOWLING & Р. 
MARSH, 2002, Wendell Lee Minckley 1935- 
2001. Copeia, 2002(1): 258-262. 

DAVIS, O. K, 1995, Robert C. Bright. American 
Association of Stratigraphic Palynologists, 
AASP Primary Records Program [unpub- 
lished manuscript, online at: http:// 
www.palynology.org/history/rcbright.html] 

DRAKE, T. E., 1968, A scientific outpost: the first 
half century of the Nantucket Maria Mitchell 
Association. Nantucket, Massachusetts, The 
Nantucket Maria Mitchell Association. 32 pp. 

DUTRO, J. T., Jr., 2004 [18 October], A national 
research laboratory in the late 20th Century: 
U.S. Geological Survey’s Paleontology and 
Stratigraphy Branch as a case study. Proceed- 
ings of the California Academy of Sciences, 
55 (Supplement 1): 177-184. 

EILPERIN, J., 2006 [14 December], USGS sci- 
entists object to stricter review rules: pre-pub- 
lication policy seen as cumbersome. 
Washington Post, December 14: p. A-29. 

HANNA, G. D., 1923 [2 January], Upper Miocene 
lacustrine mollusks from Sonoma County, Cali- 
fornia. Proceedings of the California Academy 
of Sciences, 12(3): 31-41, pls. 1-3. 

HANSELMAN, V., 1968 [20 March], The Ameri- 
can Malacological Union, Inc. Pacific Division 
Twentieth Annual Meeting [summaries]. The 
American Malacological Union, Inc., Annual 
Reports, for 1967: 61-67. 

HARRY, H. W. & B. HUBENDICK, 1964, The 
freshwater pulmonate Mollusca of Puerto Rico. 
Meddelanden fran Göteborgs Musei Zoologiska 
Avdelning, 136: 1-77 [Reprinted, Göteborgs 
Kungliga Vetenskaps- och Vitterhets-Samhälles 
Handlingar, (ser. B), 9(5): 1-77]. 

HENDERSON, J., 1931 [27 January], A new 
lymnaeid from Idaho. The Nautilus, 44(3): 75- 
TR р: © 

HERSHLER, R., 1984, The hydrobiid snails (Gas- 
tropoda: Rissoacea) of the Cuatro Ciénegas 
Basin: systematic relationships and ecology of 
a unique fauna. Journal of the Arizona-Nevada 
Academy of Science, 19(1): 61-76. 

HERSHLER, R., 1985 [9 July], Systematic revi- 
sion of the Hydrobiidae (Gastropoda: 
Rissoacea) of the Cuatro Ciénegas Basin, 
Coahuila, México. Malacologia, 26(1-2): 31- 
123. 

HERSHLER, R., 1993 [2 October], New fresh- 
water snails of the genus Pyrgulopsis 
(Rissooidea: Hydrobiidae) from California. The 
Veliger, 38(4): 343-373. 

HERSHLER, R., 1994 [19 July], A review of the 
North American freshwater snail genus Pyrgu- 
lopsis (Hydrobiidae). Smithsonian Contribu- 
tions to Zoology, 554: iv + 115 pp. 

HERSHLER, R., 2001 [18 June], Systematics of 
the North and Central American aquatic snail ge- 
nus Tryonia (Rissooidea: Hydrobiidae). Smith- 


DWIGHT WILLARD TAYLOR 217 


sonian Contributions to Zoology, 612: 53 pp. 

HERSHLER, R., 1994 [1 July], Two new genera 
of hydrobiid snails (Prosobranchia: Rissooidea) 
from the northwestern United States. The Ve- 
liger, 37(3): 221-243. 

HERSHLER, К. & T. J. FREST, 1996 [24 July], А 
review of the North American freshwater snail 
genus Fluminicola (Hydrobiidae). Smithsonian 
Contributions to Zoology, 583: iv + 41 pp. 

HERSHLER, R. 8 J. J. LANDYE, 1988 [13 April], 
Arizona Hydrobiidae (Prosobranchia: 
Rissoacea). Smithsonian Contributions to Zo- 
ology, 459: iv + 63 pp. [New taxa by Hershler 
alone]. 

HERSHLER, R. & H.-P. LIU, 2004a [2 January], 
Taxonomic reappraisal of species assigned to 
the North American freshwater gastropod sub- 
genus Natricola (Rissooidea: Hydrobiidae). 
The Veliger, 47(1): 66-81. 

HERSHLER, R. & H.-P. LIU, 2004b [1 April], A 
molecular phylogeny of aquatic gastropods 
provides a new perspective on biogeographic 
history of the Snake River Region. Molecular 
Phylogenetics and Evolution, 32: 927-937. 

HERSHLER, R., H.-P. LIU & B. K. LANG, 2007, 
Genetic and morphologic variation of the Pecos 
assiminea, an endangered mollusk of the Rio 
Grande region, United States and Mexico 
(Caenogastropoda: Rissoidea: Assimineidae). 
Hydrobiologia, 579: 317-335. 

HERSHLER, R., H.-P. LIU & С.А. STOCKWELL, 
2002 [3 April], A new genus and species of 
aquatic gastropods (Rissooidea: Hydrobiidae) 
from the North American Southwest: phyloge- 
netic relationships and biogeography. Proceed- 
ings of the Biological Society of Washington, 
115(1): 171-188. 

HERSHLER, R., H.-P. LIU & F. G THOMPSON, 
2003, Phylogenetic relationships of North 
American nymphophiline gastropods based on 
mitochondrial DNA sequences. Zoologica 
Scripta, 32(4): 357-366. 

HERSHLER AR: НР LOTE FREST EL 
JOHANNES 8 W. H. CLARK, 2006, Genetic 
structure of the western North American 
aquatic gastropod genus Taylorconcha with 
description of a second species. Journal of 
Molluscan Studies, 72(2): 167-177. 

HERSHLER, R. 8 С. LONGLEY, 1986 [14 May], 
Hadoceras taylori, a new genus and species 
of phreatic Hydrobiidae (Gastropoda: Risso- 
acea) from south-central Texas. Proceedings 
of the Biological Society of Washington, 99(1): 
121-136. 

HERSHLER, R. & G. LONGLEY, 1987 [19 May], 
Phreatoceras, a new name for Hadoceras 
Hershler and Longley, 1986 (Gastropoda) non 
Strand, 1934 (Cephalopoda). Proceedings of the 
Biological Society of Washington, 100(2): 402. 

HERSHLER, R. & Е. С. THOMPSON, 1987, 
North American Hydrobiidae (Gastropoda: 
Rissoacea): redescription and systematic re- 
lationships of Tryonia Stimpson, 1865 and 
Pyrgulopsis Call and Pilsbry, 1886. The Nauti- 
lus, 101(1): 25-32. 

HERSHLER, R. & F. G. THOMPSON, 1992, A 
review of the aquatic gastropod subfamily 


Cochliopinae (Prosobranchia: Hydrobiidae). 
Malacological Review, Supplement, 5: 1-140. 

HISLOP, S., 1860, Description of fossil shells, 
from the above-described deposits. Quarterly 
Journal of the Geological Society of London, 
16: 166-182, pls. 5-10. 

HUBENDICK, B., 1951 [22 December], Recent 
Lymnaeidae: their variation, morphology, tax- 
onomy, nomenclature, and distribution. 
Kungliga Svenska Vetenskapsakademiens 
Handlingar, 3(1): 1-223, 5 pls. 

INTERNATIONAL COMMISSION ON ZOOLOGI- 
CAL NOMENCLATURE, 1999, International 
Code of Zoological Nomenclature (4* ed.). 
London: International Trust for Zoological No- 
menclature. xxx + 306 pp. 

IOGANZEN, В. С. & Ya. |. STAROBOGATOV, 
1982, O nakhodke v Sibiri presnovodnogo 
Molliuska semeistva Triculidae (Gastropoda, 
Prosobranchia) [A finding of a freshwater mol- 
lusc of the family Triculidae (Gastropoda, 
Prosobranchia) in Siberia]. Zoologischeskii 
Zhurnal, 61(8): 1141-1147. 

JAMESON, G. & L. McMILLIN, 1985, Footprints 
on the sands of time: the writings of Dr. 
Raymond M. Alf. Claremont, California: 
Raymond M. Alf Museum of Life. iv + 120 pp. 

KABAT, A. R. & R. HERSHLER, 1993 [19 Octo- 
ber], The prosobranch snail family Hydrobiidae 
(Gastropoda: Rissooidea): review of classifi- 
cation and supraspecific taxa. Smithsonian 
Contributions to Zoology, 547: iv + 94 pp. 

KEEN, A. M. 1971, Sea Shells of Tropical West 
America: Marine Mollusks from Baja Califor- 
та to Peru. 2" ed. Stanford, Calfornia, Stanford 
University Press. xvi + 1064 pp., 22 pls. 

KORNIUSHIN, A. V. & R. K. MELNYCHENKO, 
2002 [10 October], [International Scientific 
Conference “Molluscs: The Main Results, Prob- 
lems and Perspectives of Research” 
(Zhytomyr, 13-15 Мау 2002)]. Vestnik Zoologii/ 
Journal of Schmalhausen Institute of Zoology 
[Kiev], 36(5): 91 [in Ukrainian]. 

LAMB, R. V., 1989, Nonmarine Mollusks of Pit 
91, Rancho La Brea, Southern California, and 
Their Paleoecologic and Biogeographic Impli- 
cations. Master’s Thesis, California State Uni- 
versity at Northridge. xvii + 365 pp. [not seen]. 

LIU, H.-P. & R. HERSHLER, 2005, Molecular 
systematics and evolution of western North 
American nymphophiline gastropods. Molecu- 
lar Phylogenetics and Evolution, 34: 284-298. 

LOFGREN, D., 2000, In memory of Ray. Quest: A 
Newsletter for Friends and Supporters of the 
Raymond M. Alf Museum of Paleontology, Win- 
ter/Spring 2000: 1-3 [Online at: http:// 
www.alfmuseum.org/visitorpages_ peccary.htmi]. 

LOFGREN, D., 2005, Dr. Raymond M. Alf: a Man 
Who Made the Most of His Moment in Time. 
Claremont, California, Raymond M. Alf Mu- 
seum of Paleontology. 44 pp. 

MALDE, H. E., 1972, Stratigraphy of the Glenns 
Ferry Formation from Hammett to Hagerman, 
Idaho. United States Geological Survey Bulle- 
tin, 1331D: 1-19, 2 pls. 

MALDE, H. E. & H. A. POWERS, 1962 [Octo- 
ber], Upper Cenozoic stratigraphy of Western 


218 KABAT & JOHNSON 


Snake River Plain, Idaho. Geological Society 
of America Bulletin, 73(10): 1197-1220, 1 pl. 
MARTIN, C., 1972, Know your fellow club mem- 

bers. The Festivus, 3(11): 7-13. 

MIROLLI, M., 1958 [June], | gasteropodi costieri 
del Lago Maggiore e di alcuni laghi vicini. 
Memorie dell’Istituto Italiano di Idrobiologia 
dott. Marco De Marchi, 10: 209-316. 

MORRISON, J. P. E., 1949 [April], The cave 
snails of eastern North America. The Ameri- 
can Malacological Union, News Bulletin and 
Annual Report for 1948: 13-15. 

PEARSON, J., 1995, Painfully rich: the outra- 
geous fortune and misfortunes of the heirs of 
J. Paul Getty. New York, St. Martins Press. xv 
+ 270:pp., 16 pls. 

PONDER, W. & P. BOUCHET, 2005 [20 July], 
Working classification of the Gastropoda: mod- 
ern Caenogastropoda, modern lower 
Heterobranchia. In: Р. BOUCHET & J.-P. ROCROI, 
Classification and nomenclator of gastropod 
families. Malacologia, 47(1-2): 1-397 [Hydro- 
biidae sensu lato: рр. 251-252]. 

REEDER, К. L. & В. ROTH, 1988 [3 October], A 
new subgenus of Helminthoglypta (Gas- 
tropoda: Pulmonata: Helminthoglyptidae) with 
the description of a new species from San Ber- 
nardino County, California. The Veliger, 31(3- 
4): 252-257. 

ROGERS, D. C. & A. R. WETHINGTON, 2007 
[December], Physa natricina Taylor, 1988, jun- 
ior synonym of Physa acuta Draparnaud, 1805 
(Pulmonata: Physidae). Zootaxa, 1662: 45-51. 

RONAYNE, D., 2007, Aldrich ‘Drich’ Bowler left 
a legacy in the arts. Idaho Statesman (Boise), 
January 28, p. 4. 

ROTH, B., 1986, Land mollusks (Gastropoda: 
Pulmonata) from Early Tertiary Bozeman 
Group, Montana. Proceedings of the Califor- 
nia Academy of Sciences, 44(11): 237-267. 

ROTH, B., 2003 [6 October], Introduction to 
Physidae (Gastropoda: Hygrophila); biogeog- 
raphy, classification, morphology [Review of 
Taylor, 2003]. The Veliger, 46(4): 364-365. 

RUSSELL, R. H., 1971 [31 December], Mollusca 
of Fish Springs, Juab County, Utah: rediscov- 
ery of Stagnicola pilsbryi (Hemphill, 1890). 
Great Basin Naturalist, 31(4): 223-236. 

SMITH, С. R., Т.Е. DOWLING, К. W. GOBALET, 
T. LUGASKI, D. K. SHIOZAWA & R. P. EVANS, 
2002 [11 December], Biogeography and tim- 
ing of evolutionary events among Great Basin 
fishes. Smithsonian Contributions to the Earth 
Sciences, 33: 175-234. 

SMITH, J. T., 1978 [14 March], Primary types in 
the Stanford paleontological type collection. 
Bulletins of American Paleontology, 72(300): 
317-552. 

STAROBOGATOV, YA. |., 1967 [after 25 Octo- 
ber], K postroeniiu sistemy presnovodnykh 
legochnykh molliuskov [On the systematization 
of freshwater pulmonate molluscs]. Trudy 
Zoologicheskogo Instituta, 42: 280-304. 

STAROBOGATOV, Ya. |., 1970 [after 15 Octo- 
ber], Fauna Molliuskov i Zoogeograficheskoe 
Raionirovanie Kontinental’nykh Vodoemov 


Zemnogo Shara [Fauna of molluscs and zoo- 
geographical separation into districts of the 
continental water reservoirs of the world]. 
Leningrad, Zoologicheskii Institut, Akademiiia 
Nauk SSSR. 372 pp. 

TE, С. A., 1975, Michigan Physidae, with sys- 
tematic notes on Physella and Physodon 
(Basommatophora: Pulmonata). Malacological 
Review, 8: 7-30. 

TE, С.А., 1980 [25 April], New classification sys- 
tem for the family Physidae (Pulmonata: 
Basommatophora). Archiv für Molluskenkunde, 
110(4-6): 179-184. 

THIELE, J., 1928 [12 September], Revision des 
Systems der Hydrobiiden und Melaniiden. 
Zoologische Jahrbücher, Abteilung für 
Systematik, Okologie und Geographie der 
Tiere, 55(5-6): 351-402, pl. 8. 

THOMPSON, F. G., 1979, The systematic status 
of the hydrobioid snail genus Nymphophilus 
Taylor 1966 and the status of the subfamily 
Nymphophilinae. Malacological Review, 12(1- 
2): 41-49. 

TRACEY, S., J. А. TODD & D. H. ERWIN, 1993, 
Mollusca: Gastropoda. Pp. 131-168, in: M. J. 
BENTON, ed., The Fossil Record 2. London, 
Chapman Hall. xvii + 845 pp. [Note: the bibli- 
ography erroneously attributes several papers 
by John D. Taylor to Dwight W. Taylor]. 

TRYON, G. W., 1866 [1 April], [Review of “Re- 
searches upon the Hydrobiinae and Allied 
Forms,” by W. Stimpson (1865).] American 
Journal of Conchology, 2(2): 152-158. 

WALLACE, R. & M. P. CARR, 1986, The Willard 
Hotel: an illustrated history. Washington, D.C., 
Dicmar Publishing. 112 pp. 

WEBB SCHOOL, 1949, E/ Espejo [yearbook]. 
Claremont (California). 

WETHINGTON, A. R. & C. LYDEARD, 2007, A 
molecular phylogeny of Physidae (Gastropoda: 
Basommatophora) based on mitochondrial 
DNA sequences. Journal of Molluscan Stud- 
les, 73(3): 241-257. 

WEST, R., 1973, Getty son’s death laid to over- 
dose of alcohol, drugs. Los Angeles Times, 
June 8, 1973, p. A-3. 

WILKE, T., G. M. DAVIS, A. FALNIOWSKI, F. 
GIUSTI, M. BODON & M. SZAROWSKA, 2001 
[31 December], Molecular systematics of 
Hydrobiidae (Mollusca: Gastropoda: Ris- 
sooidea): testing monophyly and phylogenetic 
relationships. Proceedings of the Academy of 
Natural Sciences of Philadelphia, 151: 1-21. 

WOO, E., 1999, Raymond Alf: teacher at Webb 
Schools, founder of paleontology museum. Los 
Angeles Times, October 2, p. A-20. 

WU, S.-K. & N. E. BRANDAUER, 1982 [1 Octo- 
ber], Type specimens of Recent Mollusca in 
the University of Colorado Museum. Natural 
History Inventory of Colorado, 7: 1-47. 

YEN, T.-C., 1946 [September], Late Tertiary 
fresh-water mollusks from southeastern Idaho. 
Journal of Paleontology, 20(5): 485-494, pl. 76. 


Revised ms. accepted 13 December 2007 


MALACOLOGIA, 2008, 50(1-2): 219-264 


THE MOLLUSCAN TAXA MADE AVAILABLE IN THE 
GRIFFITH & PIDGEON (1833-1834) EDITION OF CUVIER, 
WITH NOTES ON THE EDITIONS OF CUVIER AND ON WOOD'S 
INDEX TESTACEOLOGICUS 


Richard E. Petit? 8 Eugene V. Coan?* 


ABSTRACT 


This paper reviews the new taxa made available by Gray in the Griffith 8 Pidgeon (1833- 
1834) English translation of Cuvier's famous Le regne animal (1830), as well as the taxa 
attributed, correctly or incorrectly, to Gray in this work. We discuss various complications 
concerning the dating and authorship of these taxa, and some new information is provided 


about their type material. 


Six generic names date from this work, five of them now considered valid. Sixty-five 
species names were made available, of which 44 are now considered valid, although 
some of these have not been recognized in recent literature, and one is a nomen dubium; 
21 are placed in synonymy because they are junior synonyms, preoccupied homonyms, or 
unused senior synonyms that have been or should be suppressed. 

Key words: taxonomy, nomenclature, Griffith, Pidgeon, J. E. Gray, W. Wood, Cuvier, 


Mollusca, bibliography. 


INTRODUCTION 


The Mollusca volume (Volume 12, 1833- 
1834) of Griffith 8 Pidgeon’s series of transla- 
tions of Cuvier's Le Regne Animal (1830) is in 
part a translation of the molluscan volume of 
Cuvier’s second edition. It has long been 
known as a source of new taxa. The new gen- 
era and species, attributed either to Gray or to 
Griffith & Pidgeon, ex Gray ms, have mostly 
been incorrectly dated as 1834. Some new taxa 
in this work have been missed altogether be- 
cause they were misattributed to other authors. 

We have reviewed the text of this important 
work, weighed the evidence as to the author- 
ship and dates of its new taxa, made a prelimi- 
nary effort to locate the type material of the 
new species, and explored the taxonomic con- 
sequences of the resulting conclusions. Some 
of the taxa in this work that have been 
misattributed are discussed, and an annotated 
bibliography of several related editions of 
Cuvier’s seminal work is appended. 


Historical Context 


Georges Cuvier (b. 1769 — d. 1832), after 
having published individual anatomical stud- 


ies on many animals, including a number of 
species of mollusks, produced the first edition 
of a comprehensive overview of animal life 
(Cuvier, 1816). This was a four-volume set, 
with the Mollusca, in the modern sense, rep- 
resented in volume 2: 351—494; the only fig- 
ures of mollusks were on a single plate 
included in volume 4 (plate 11). For Cuvier and 
others of his time, the Mollusca also included 
the tunicates, brachiopods, and barnacles. 
Smith (1993) provides an exhaustive bibliog- 
raphy of Cuvier’s publications. 

Later, Cuvier (1829-1830), in collaboration 
with Pierre-Andre Latreille (b. 1762 — а. 1833), 
produced a five-volume second edition of Le 
Règne Animal, augmented by recent discov- 
eries, notably those made by naturalists trav- 
eling with French expeditions around the 
world. The coverage of the Mollusca in the 
modern sense, now in volume 3, was ex- 
panded from 144 pages in 1816 to 162 pages 
in 1830. No new figures were added, with the 
same plate, now numbered 14, included with 
the related text in volume 3. 

Beginning in 1824, the English biologists Ed- 
ward Griffith (b. 1790 — 4. 1858) and Edward 
Pidgeon (b. 1790? — d. 1834) began an En- 
glish version of Cuvier. Gruber (2004) noted that 


1806 St. Charles Rd., North Myrtle Beach, South Carolina, U.S.A. 
“Research Associate, Santa Barbara Museum of Natural History, 2559 Puesta del Sol Rd., Santa Barbara, California, U.S.A. 


*Corresponding author: gene.coan@sierraclub.org 


220 PETIT & COAN 


“Although Griffith was the controlling editor and 
a contributing author (for instance on fishes), 
the major part of the translation was done by 
Pidgeon who also produced the volume on fos- 
sils.” Gruber termed Pidgeon a “working part- 
ner” and listed John E. Gray as one of three 
“specialist associates.” The work eventually 
amounted to 16 volumes and was completed 
in 1835. Their labor became complicated when 
Cuvier’s second edition started to appear after 
they had begun translating the first, and their 
translation had to switch editions in 1829. 
Cuvier had also published, in 1826, the third 
edition of his Recherches sur les Ossements 
Fossiles that Pidgeon had been working on in- 
dependently. Pidgeon’s translation of that work 
was introduced into the series as Volume 11, 
the volume number usually being placed in 
brackets to indicate, among other things, that 
itis not part of the translation of Cuvier’s Regne 
Animal. The Mollusca were contained in Griffith 
& Pidgeon’s volume 12, published after 
Cuvier’s second edition was completed, and 
their translation is thus based on that edition. 

Pages 1-138 of Griffith & Pidgeon are a trans- 
lation of the “Mollusca” of Cuvier (1830), with 
the true Mollusca covered on pages 1-125, 
pages 125-138 being a translation of Cuvier’s 
text on tunicates, brachiopods, and barnacles. 
In their translation, Griffith & Pidgeon shortened 
Cuvier’s footnotes, incorporating the abbrevi- 
ated versions of them into their main text. Their 
work on the Mollusca, however, was not lim- 
ited to a mere translation of Cuvier’s second 
edition. Pages 139—413 constitute а Supple- 
ment to the Mollusca, which seems to have 
been entirely original; pages 413—434 supple- 
ment coverage of the tunicates, brachiopods, 
and barnacles. Pages 435-522 of the volume 
contain a translation of Cuvier’s Zoophytes, 
essentially the rest of the non-arthropod inver- 
tebrates, and pages 523-594 constitute Griffith 
& Pidgeon’s Supplement to those. 

While the Supplementary treatment of the 
Mollusca is a comprehensive and engaging 
review of the literature, it is of little taxonomic 
significance, although it does contain a few 
misspelled generic names and newly or mis- 
takenly Latinized terms (Appendix A). The 
absence of any reference in the text to the 
plates is puzzling as they could have been 
referenced to advantage in appropriate places. 

Finally, pages 595-601 contain an Index to 
the plates on the “Mollusca” in the broad sense 
of Cuvier, discussed further below. There is 
no such index for the plates on the Zoophytes. 


Two Reviews 


An uncomplimentary contemporary review of 
Griffith & Pidgeon’s Mollusca volume was writ- 
ten by Baron André Etienne Justin Pascal Jo- 
seph François d’Audebard de Férussac (b. 
1786 — d. 1836) in 1835. Part of that review 
commenting unfavorably on the plates will be 
detailed below. In his closing paragraph, here 
translated loosely, Ferussac commented that 
in a work of this type each part should be di- 
rected by a recognized specialist, and Mr. 
Griffith's The Animal Kingdom appeared to 
miss that goal as the various classes were not 
always directed by capable specialists. 

A long review was also published by Rudolph 
Amandus Philippi (b. 1808 — а. 1904) in 1848, 
a rather late appearance for such a detailed 
review. Philippi listed the new taxa by name, 
plate and figure number, with comments about 
many, also noting errors. With six exceptions, 
Philippi attributed authorship of the new spe- 
cies to Gray. Those six are attributed to [Griff?] 
(square brackets and query as in Philippi’s 
original) without any stated reason. However, 
these six names were listed without authors 
in Griffith & Pidgeon’s Index, Gray’s name 
having been inadvertently omitted. As will be 
discussed below, the names date from the 
plates. 

Philippi inserted a footnote listing nine ep- 
onyms among the new taxa, stating that Gray 
perpetuated in this work good friends. The nine 
listed are all females although that obvious 
point was not mentioned by Philippi. The male 
eponyms introduced by Gray were not singled 
out for special notice. 


The Plates 


To further enhance their treatment of the 
Mollusca, represented in Cuvier’s two editions 
by the same rather crude plate, Griffith and 
Pidgeon included 41 Mollusca plates, none 
that of Cuvier. There is one plate of barnacles 
(plate 2); two of brachiopods (plates 6, 7); and 
two of tunicates (plates 9, 10). The other 36 
plates contain true mollusks. There are also 
20 separately numbered plates to accompany 
the treatment of the Zoophytes. Each plate has 
a legend at the bottom with species names 
but not authors. (Here we use the term “leg- 
end” to apply to information that appears on a 
plate itself about the figures thereon, the term 
“caption” being restricted to similar informa- 
tion that appears on another page.) 


MOLLUSCAN TAXA IN GRIFFITH 8 PIDGEON (1833-1834) 221 


The figures on the molluscan plates were 
partly obtained from the /conographie (1829— 
1844) of Félix-Edouard Guérin-Méneville (b. 
1799 —d. 1874), originally produced to accom- 
pany and illustrate Cuvier’s second edition. 
Unfortunately, the dates of the 38 beautiful, 
accurate Guérin-Méneville plates and their 
legends remain unknown. It is known that one 
mollusk plate appeared in 1829, four more in 
1830, and two more in 1832, and all had ap- 
peared by 1837, but it is not known which ap- 
peared when; the explanatory text was 
published in 1844 (Cowan, 1971). (Appendix 
C herein provides additional information about 
this work.) In Griffith & Pidgeon, Mollusca 
(sensu lato) plates 2-12, 27, 32, 33, 35, 38, 
and 39, none of which contains new taxa, are 
copied entirely or in substantial part from 
Guérin-Méneville. Of these 17 plates, 12 are 
copied in their entirety with figures unchanged 
in size; the other five — plates 27, 32, 33, 38, 
39 — are each wholly, or in part, composed of 
figures from more than one Guérin-Méneville 
plate, most figures considerably reduced in 
size. These plates are generally good copies, 
but there are exceptions, and it is thus unlikely 
that the copy work was all done by one arti- 
san. 

Mollusca plates 15-18 of Griffith & Pidgeon 
were copied from the Manuel de Malacologie 
et de Conchyliologie (1825-1827) of Henri 
Marie Ducrotay de Blainville (b. 1777 - d. 
1850). About these plates, designed to illus- 
trate shell morphology, Griffith & Pidgeon (p. 
280) stated: 

“As we have inserted four plates in explana- 
tion of the terms made use of in conchology, it 
becomes necessary to advert here to such of 
the references as could not be engraved on 
the coppers.” 

That statement is followed by two pages of 
descriptive terminology. It is notable that this 
is the only mention of any of the plates, or the 
figured specimens, made in the Griffith & 
Pidgeon text. 

The other Mollusca plates — 1, 13, 14, 19- 
26, 28-31, 34, 36, 37, 40, and 41 — containing 
mollusks in the modern sense were new, and 
their contents were supplied by the British 
Museum under the direction of John Edward 
Gray (b. 1800 — d. 1875). Plate 14 includes 
the legend “W. Hawkins, ае!.” (Benjamin 
Waterhouse Hawkins, b. 1807 —d. 1889). The 
names of the artist(s) and engraver(s) are not 
shown on any other plates. Judging from the 
shading within the gastropod apertures, it is 


probable that more than one artist and/or en- 
graver was involved. Of the 14 original plates 
of gastropods, four are drawn in the “French 
fashion”, with the apex at the bottom and the 
aperture at the top. 

The majority of the 20 Zoophyte plates were 
also copied from Guérin-Méneville. The ab- 
sence of an Index to Zoophyte plates or any 
other mention of these plates, and their fig- 
ures is inexplicable. 

The plates, as well as the work in its entirety, 
were denigrated by Férussac (1835) in his 
review, and he stated that while he had praised 
the beauty of the plates in the first volumes of 
Griffith & Pidgeon in an earlier review 
(Férussac, 1825), he regretted that the plates 
of volume 12 are poorly drawn and engraved. 
He stated that the plates were so defective as 
to often make identification of the species very 
difficult. As usual in a work with plates intended 
to be colored, the uncolored plates are not par- 
ticularly attractive. However, in our opinion, 
neither version is as bad as claimed by 
Férussac. Férussac listed a number of non- 
marine species with comments on their validity - 
Or supposed synonymy and stated that “all 
these species, especially the helices, are al- 
most unrecognizable” [Toutes ces espéces, 
surtout les hélices, sont presque mécon- 
naissables]. Ferussac’s comments about the 
utility of the plates are not warranted and evi- 
dently some unstated agenda or prejudice af- 
fected his review. He certainly had a “proprietary 
interest” in the helicids. 

Férussac commented on the fact that a num- 
ber of plates had been copied, but erred in 
the origin of some. He said that one was taken 
from his own work (which work not stated), 
but the plate number specified is one copied 
from Guérin-Méneville. He also made much 
of the fact that plates 28 and 36 were engraved 
in mirror-image, stressing the fact that they had 
not only been so engraved, but “enluminées 
et distribuées aux souscripteurs” [emphasis in 
Original]. He somehow overlooked the fact that 
the figures on plate 37 were also reversed, 
possibly as that plate illustrates marine spe- 
cies, and Férussac's review mentioned only 
non-marine taxa. These three plates are dis- 
cussed below under the dating of this work. 

Férussac also expressed surprise that for a 
project that had started so well, and was sev- 
eral years in progress, that the authors lacked 
the self-esteem to offer only original figures. 
This ignored the fact that plates from Guérin- 
Méneville had been copied in other volumes. 


222 PETIT & COAN 


Although the species listings in the plate leg- 
ends do not provide the names of the authors 
of the contained species, the Index to the 
plates does list most. In the Index, many of 
the taxa, including the new species, are at- 
tributed to Gray. 


Authorship of the New Taxa 


The authorship of the new taxa contained in 
Griffith & Pidgeon has been the source of some 
confusion, particularly in recent years. Our in- 
terpretation of authorship under provisions of 
the ICZN Code Art. 50.1 relies upon several 
factors, both internal and external. The Code 
requires “that if it is clear from the contents 
that some person other than the author is alone 
responsible for both the name or act and for 
satisfying the criteria of availability other than 
actual publication, then that other person is 
the author of the name or act.” 

(1) Griffith and Pidgeon did not engage in 
taxonomic work on the Mollusca, in this work 
or elsewhere, and their Supplementary text on 
the Mollusca covers morphology, physiology, 
and natural history, not taxonomy or nomen- 
clature. No new taxa were attributed to either 
of them in the Index. 

(2) A footnote on the first page of the Index 
(p. 595) stated that “most of the inedited shells 
figured in this work are from the collection in 
the British Museum,” the term “inedited” mean- 
ing not previously published. The Mollusca in 
the British Museum were then, of course, un- 
der the care of John Edward Gray. Griffith & 
Pidgeon were not only well acquainted with 
Gray but appreciated his expertise, as evi- 
denced in their Supplement on Mollusca (p. 
154): “this branch of zoology has been much 
indebted ... to some distinguished men 
amongst ourselves, of whom we shall merely 
mention Dr. Leach and Mr. Gray ...” 

(3) The entirety of plates 1, 13, 14, 19-26, 
28—31, 34, 36, 37, 40, and 41 seem to have 
been based on specimens then in the British 
Museum, are all of similar style, and either the 
specimens or the drawings used to engrave 
them were thus probably supplied by Gray. 
This set of plates thus constitutes a discrete 
“unit” within the volume. 

(4) One of the new taxa, Clavatula griffithii, 
is dedicated to Griffith. 

(5) Gray contributed to other volumes in the 
series, such as in the insects (Cowan, 1969: 
140). According to Gunther (1975: 173): “Justly, 


his [Gray’s] name is associated with Cuvier 
in Griffith's English translation of the Règne 
Animal (1827-1830) in which he was respon- 
sible for contributions to mammals, reptiles, 
birds and, unacknowledged, for the Mol- 
lusca.” 

(6) To the extent that type material has thus 
far been located in The Natural History Mu- 
seum in London, it was labeled “Gray” or “Gray, 
Griffith & Pidgeon” and catalogued as such. 

(7) Almost all subsequent treatments of the 
new taxa made available in Griffith & Pidgeon 
attribute them to Gray, particularly those in the 
decades immediately subsequent to their pub- 
lication. This includes those of Gray himself. 
For example, in a footnote in the list of the 
Volutidae in the British Museum (Gray, 1855: 
5), he wrote: “In 1833 | described and figured 
three species of this genus in the Mollusca 
plates to Griffith’s translation of Cuvier’s ‘Ani- 
mal Kingdom’ under the names Melo miltonis, 
M. georginae and M. Broderipii ...”. The new 
names were also so treated by Sherborn 
(1922-1932) and Neave (1939-1940). 

In recent years, most workers continue to 
favor this interpretation. We thus believe it is 
in the interests of stability to continue this con- 
sensus, if necessary with an application to the 
International Commission on Zoological No- 
menclature. 

While Gray seems to have supplied some 
descriptive and locality notes covering his 
plates for the Index, including some updated 
nomenclatural information, that Index also in- 
cludes the plates obtained from the other 
sources, and it contains many errors and 
misattributions of species. This is considered 
evidence that someone other than Gray pre- 
pared the Index. One entry on p. 597 of the 
Index provides evidence that its preparer(s) 
regarded Gray as the author of the new taxa: 
under Cyllene owenii Gray, it is stated that “Mr. 
Gray has, therefore, separated it generically.” 

That the actual production of the work, plates 
and Index, was not in Gray’s hands is evinced 
by printing problems with the three reversed 
plates, as well as by the many errors in the 
Index. 

Wood's Index testaceologicus (1828b) must 
here be introduced into this narrative because 
many names that first appeared in this work 
have also been credited to Gray by subsequent 
workers. As William Wood (b 1774 — 4. 1857) 
did not indicate authorship of any species, no 
credit for any contribution Gray made is to be 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 225 


found among the new species names. How- 
ever, in his Preface, Wood (1828b: iii) “grate- 
fully acknowledges the assistance of Mrs. 
Mawe and Mr. Gray, from whose Cabinets, and 
principally from the former he has derived the 
most essential benefit.” In the lists of taxa, 
where the new names appear, the abbrevia- 
tion “M. Cab.” is used for specimens from Mrs. 
Mawe’s Cabinet, “Gray Cab.” for Mr. Gray’s 
Cabinet, and “Br. M.” for British Museum. 
Dance (1972) discussed the various editions 
of Wood, stating that Hanley (1856) “made 
matters worse by attributing authorship to Gray, 
Mawe or Leach rather than to Wood.” The 
matter of authorship was also touched on by 
Wilkins (1957: 157-158), who warned that 
even after the advent of Sherborn’s Index 
Animalium, “caution is still necessary when 
dealing with the combined efforts of Gray and 
Wood.” 

Gray’s contribution to Wood’s work was also 
discussed by Beu & Rooij-Schuiling (1982: 
215), who suggested that “perhaps a case 
could even be made for regarding Gray as the 
author of new names in this book [Wood, 
1828b]”. We are not prepared to go that far, 
although some of the names based on Gray’s 
material, or that of the British Museum, were 
elsewhere attributed to Gray by others, as well 
as by Gray himself. It must be remembered 
that not until the latter half of the 19" century 
was there a generally accepted Code of No- 
menclature. The early Codes did not address 
authorship, which was generally attributed to 
the person deemed responsible for providing 
the name, not necessarily the person publish- 
ing it. Unlike the Griffith & Pidgeon work, there 
are no discrete pages or plates in Wood that 
can be attributed to Gray alone, and so, un- 
-der the existing Code, Wood is the author. If it 
is so desired, optional credit can be supplied 
as “ex Gray ms” in appropriate cases. 

Because of their unusual format, Wood’s 
works (1828a, b) can be a source of confu- 
sion for authors who have not worked with 
them extensively. In Appendix B, we discuss 
their format in detail. 


Dating Griffith & Pidgeon 


Cowan (1969) produced the only available 
collation of Griffith & Pidgeon, covering all 16 
volumes of the series. Cowan concluded that 
volume 12 appeared as parts 38 (December 
1833), 39 (March 1834), and 40 (June 1834), 
based on an assumed three-month interval for 


issuance of the parts. The same part numbers 
were listed by Ferussac (1835: 73) without 
dates. Unfortunately, the pagination of the 
parts is not known, Cowan stating (p. 138) that: 
“Pagination has not been attempted at this 
stage. There are several indications that 192 
pages were arrived at as the standard for each 
part. The plates were probably published 
bound into their respective parts.” 

The total pagination for the volume is viii + 
601 pp., 41 + 20 pls. Page 192 is the end of a 
signature, as is page 384. With a total of 601 
(+ viii) pages, the figure of 192 pages cannot 
quite hold for the third part, but it is close. Page 
592 is the end of a signature. Page 593 is the 
beginning of signature Qq, and page 601 bears 
the signature letters Rr. Because signature Qq 
is thus only 8 pages (a half-signature), it is 
probable that signature Qq also contained 
pages i-viii, and page 601 was printed as a 
single page. 

Cowan’s suggestion that the plates were 
printed with their respective parts in which they 
were to be bound does not match their place- 
ment. Indeed, their arrangement is rather hap- 
hazard, and plates can be located within the 
bound volume only by referring to the List of 
Plates on pages vii-viii, which indicates where 
in the bound text each is to be placed. The 
majority of the plates are placed in the same 
general area as the related text. However, 36 
Mollusca (in Cuvier’s broader sense) plates 
were to be bound within pages 1-192, 4 Mol- 
lusca and 2 Zoophyte plates were to be bound 
within pages 193-384, and 1 Mollusca plate 
and 18 Zoophyte plates were to be bound 
within pages 385-592. As all but two of the 
original Mollusca plates — plates 40 and 41 — 
are dated 1833, we must either accept that as 
their issue date or accept 1834 for those bound 
in subsequent to p. 192. The only plate with 
new taxa affected would be Mollusca Plate 22, 
which was to be bound opposite p. 420 (in the 
beginning of the brachiopod text). Mollusca PI. 
41, which contains new taxa, is bound as the 
frontispiece. It does not appear on the List of 
Plates, and is assumed to have been issued 
with the last signature, which included the title 
page, introduction, and final pages. 

Of the Zoophyte plates, only plate 1 is dated 
1833; Zoophyte plates 2-20 are all dated 1834. 

As further evidence that the plates appeared 
in at least two batches is the fact that three 
plates were printed with reversed images but 
correct legends. These original plates, num- 
bers 28, 36, and 37, are each dated 1833 


224 PETIT 8 COAN 


г 
= 


restora articulada. 


2 Melee argellaveer. 
3 Cyelestema pulelira 
+ Orelestona Madagascartensis. 


5lydostema auriculares. 


PRO LE ERA 


er 


Anden, Раде, bp WE 
2 


FIG. 1. Original Plate 28 from Griffith 8 Pidgeon (1833) with reversed images. Courtesy 
The Australian Museum. 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 229 


$ Ней mera. 

2 Helix virtdis Desm. 
8 Faludina: subeostata. 
+ Helix Lay: 

Helix радия. 


Gelée Frasert. 


$ ; IO 
LOR BH: вый 


FIG. 2. Original Plate 36 from Griffith & Pidgeon (1833) with reversed images. Courtesy 
The Australian Museum. 


226 PETIT & COAN 


L Lolumbellu 


AVERY, b Triton 


STO) VELA AAA 


2 Lolo herpateormeis. 


O Cfeavetiula СЕНЕ. 


Altvar tersalata yar. 7 Bulimes auris- vulpina: 


+ Prrula: striatac 8 Bulla: semaplicatas 


D Aneti¢ 


LOAN Published: by Whittaker 202, 


Tre SHAS 


FIG. 3. Original Plate 37 from Griffith & Pidgeon (1833) with reversed images. Courtesy 
The Australian Museum. 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 227 


TABLE 1. Most likely dates of volume 12 of Griffith & Pidgeon ([1833]-1834). 


Part Pages Plates Date 
38 1-192 Mollusca pls. 1-27, 29-35, 38, 39 [original pls. 28, Dec. 1833 
36, 37 with reversed images] 
Zoophyte pl. 1 
39 193-384 ESE March 1834 
40 viii + 385-601 Zoophyte pls. 2-20 June 1834 


Mollusca corrected pls. 28*, 36*, 37*, pls. 40, 41 


(Figs. 1-3). When the error was discovered, 
subscribers were instructed to discard them 
and replace them with corrected versions, 
each then dated 1834. The only volume con- 
taining the reversed originals that we have lo- 
cated is the one referred to by Iredale (1937) 
in the Australian Museum in Sidney. Only two 
of them were noted by Férussac (1835) in his 
review. Oddly, Gray changed a single species 
name between the two versions of plate 36 
(see under Helix hayii below). 

Gray (1855: 5) was quite specific about hav- 
ing named the three species of Melo in 1833. 
These species appeared on Plates 26, 29, and 
34. 

While there is no way to know which of the 
two parts issued in 1834 contained the plates 
bearing that date, the most likely dates of vol- 
ume 12 of Griffith & Pidgeon are given in 
Table 1. 


Format 


In the following list, the taxa made available 
by Gray in Griffith & Pidgeon, or by Griffith & 
Pidgeon alone, are given in the headers. The 
taxa have been arranged using modern fam- 
ily grouping. If the taxa were first made avail- 
able by Gray or others in earlier publications, 
those original combinations form the headers. 

The chresonymies that follow are not in- 
tended to be exhaustive and contain those 
references needed to understand the taxon, 
its chief synonyms, and current allocation. 

Later homonyms are indicated in brackets; 
earlier homonyms are given without brackets. 

In all cases in which we refer to taxa intro- 
duced by Linnaeus, we here use the spelling 
Linnaean. In quotations, we use the orthogra- 
phy of the original. 

The Literature Cited covers all works and 
taxa mentioned. 


March or June 1834 


TAXA ATTRIBUTABLE TO GRAY IN 
GRIFFITH 8 PIDGEON, WITH NOTES 
ON OTHER RELEVANT TAXA 


GENERA 


Bullia Gray in Griffith & Pidgeon, 1833; 
Bullaea Gray — Griffith & Pidgeon, 1834 


Bullia Gray in Griffith & Pidgeon, 1833: pl. 37, 
fig. 8 [legend; first issue of pl. with reversed 
images]; 1834: pl. 37*, fig. 8 [legend; sec- 
ond issue of pl. with corrected images]. 

Bullaea Gray — Griffith & Pidgeon, 1834: 596, 
spelling error for Bullia Gray in Griffith & 
Pidgeon, 1833. 

Non Bullaea Lamarck, 1801: 63 [Gastropoda]. 

Bullia Gray, 1839: 125 [again indicated as 
being new]. 


Remarks 

Type species: Bullia semiplicata Gray in 
Griffith & Pidgeon, 1833, by monotypy. Allmon 
(1990: 19) listed this in error as being type 
species by “original designation’. Bullia is a 
valid genus of Nassariidae. See also under 
Bullia semiplicata below. The misspelling 
Bullaea was not picked up in nomenclators but 
was noted by Allmon (1990: 13). 


Glycymeris Gray in Griffith & Pidgeon, 1833; 
Glycemeris Griffith & Pidgeon, 1834 


Glycymeris Gray in Griffith & Pidgeon, 1833: 
pl. 31, fig. 2 [legend], non da Costa, 1778, 
misspelling of G/ycimeris Lamarck, 1799. 

Glycemeris — Griffith & Pidgeon, 1834: 597 
[footnote]. 


Remarks 
The Glycymeris of Gray in Griffith & Pidgeon 
(1833) was a misspelling of Glycimeris 


228 PETIT & COAN 


Lamarck, 1799: 83, an older but now sup- 
pressed senior synonym of Panopea Menard 
de la Groye, 1807: 135 (ICZN Opinion 1414, 
1986) [Hiatellidae], not G/ycymeris of da Costa, 
1778: 168, the nominal genus of the Glycyme- 
rididae. Vokes (1980: 180) interpreted Glycyme- 
ris “Griffith & Pidgeon” as the proposal of a new, 
homonymous genus and synonym of Glaucono- 
me. Spelled as Glycemeris in footnote on p. 597. 
See also under Glycymeris apinensis below. 


Cyllene 
Gray in Griffith & Pidgeon, 1834 


Cyllene Gray in Griffith & Pidgeon, 1834: pl. 
41, fig. 2 [legend]; 597. Type species by 
monotypy: Cyllene owenii Gray in Griffith & 
Pidgeon, 1834. 

Cyllene Gray — Griffith & Pidgeon, 1834: 597. 

Cyllene Gray — Gray, 1839: 108. 


Remarks 

À valid genus of the Nassariidae (Cerno- 
horsky, 1975: 166-167). See also under 
Cyllene owenii below. 


Littoraria 
Gray in Griffith & Pidgeon, 1833 


Littoraria Gray in Griffith & Pidgeon, 1833: pl. 
1, fig. 3 [legend]. 
Littoraria Gray — Griffith & Pidgeon, 1834: 598. 


Remarks 

À valid genus ofthe Littorinidae (Reid, 1986: 
71-73). Type species, by monotypy: “Littoraria 
pulchra Gray”, = Littorina pulchra G. B. 
Sowerby |, 1832a, = Turbo zebra Donovan, 
1825. See also under Littorina pulchra below. 


Neaera 
Gray in Griffith & Pidgeon, 1833 


Neaera Gray in Griffith & Pidgeon, 1833: pl. 
22, fig. 5 [legend]. 
Non Neaera Robineau-Desvoidy, 1830: 84 
[Diptera]. 

Neroea Gray — Griffith & Pidgeon, 1834: 598, 
misspelling. 

Naeera, Neraea, Neara, misspellings of later 
authors. 


Remarks 

Synonym of Cuspidaria Nardo, 1840 (Coan 
et al., 2000: 547) [Cuspidariidae]. Type spe- 
cies, by monotypy: Neaera chinensis Gray in 
Griffith 8 Pidgeon, 1833; see also under this 
species below. 


Pusio 
Gray in Griffith & Pidgeon, 1833 


Pusio Gray in Griffith & Pidgeon, 1833: pl. 25, 
figs. 1, 2. Type species, by subsequent des- 
ignation of Gray (1847: 133): Triton (Pusio) 
elegans Gray in Griffith 4 Pidgeon, 1833. 


Remarks 

Two species were originally included in this 
new subgenus, so a subsequent designation 
is required, that of Gray (1847) being the first. 
Valid genus of Buccinidae. Type species des- 
ignation is not by monotypy, as stated by 
Vermeij (2006: 86-87) and others. See also 
under Triton elegans below. 


Villorita 
Gray in Griffith & Pidgeon, 1833 


Villorita Gray in Griffith & Pidgeon, 1833: pl. 
31, fig. 5 [legend]; 1834: 601. 
Villorita Gray — Griffith 8 Pidgeon, 1834: 601. 


Type species, by monotypy: Cyrena cyprino- 
ides Gray, 1825: 136. 


Remarks 

Spelled as Velorita by Gray (1840a: 149; 
1840b: 134; 1842: 75, 91; 1847: 184). A valid 
genus of the Corbiculidae (Keen 8 Casey, 
1969: 668; Glaubrecht et al., 2007: 268). 


SPECIES 


In the following list, the phrase “Type mate- 
rial not found in 2006 in the BMNH?” indicates 
that only a superficial search could be made, 
and it is possible that the type material may 
still be there. On the other hand, the phrase 
“Type material not present in the ВММН” indi- 
cates that the particular family has been 
curated to the point at which it is unlikely that 
the type material is present. 


Bivalvia 
Corbiculidae 


Venus similis 
W. Wood, 1828 


Venus similis W. Wood, 1828b: 5, pl. 2, Venus 
TG. 3. 

Cyrena similis Gray — Griffith & Pidgeon, 1834: 
597. 

Venus similis Gray in Wood — Deshayes, 1855: 
225; 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 229 


Cyrena similis Gray in Griff. — Deshayes, 1855: 225. 
С. similis Gray — Hanley, 1856: 203, Suppl. pl. 
2, Venus fig. 5. 


Remarks 

Credited to Gray in the list of figures in Griffith 
& Pidgeon (1834: 597), Gray was perhaps re- 
sponsible for the ms name, which was first 
made available by Wood (1828b), who listed 
the “В.М.” as his source. Deshayes (1855: 225) 
listed as a Corbicula and did not show author- 
ship in any of his species headers; we have 
listed both items from his chresonymy to show 
that he also attributed the name to Gray. Local- 
ity given as China by Wood, and as “China?” 
by Deshayes (1855: 225) and Counts (1991: 
33). Type material not found in 2006 in BMNH 
but evidently there in 1855, Deshayes indicat- 
ing that the species was in the “ВМ”. Probably 
belonging in the genus Geloina Gray, 1842: 75. 


Unionidae 


Unio douglasiae 
Gray in Griffith & Pidgeon, 1833 


Unio douglasiae Gray in Griffith & Pidgeon, 
1833: pl. 21, fig. 2 [legend]. 

U. douglasiae Gray — Griffith & Pidgeon, 1834: 601. 

U. douglasiae douglasiae Gray in Griffith & 
Pidgeon, 1834 — Haas, 1969: 56-57. 


Remarks 
China (Haas, 1969). Type material not found 
in 2006 in the BMNH. 


Unio leaii 
Gray in Griffith & Pidgeon, 1833 


Unio Геай Gray in Griffith & Pidgeon, 1833: pl. 
21, fig. 1 [legend]. 

U. leaii Gray — Griffith & Pidgeon, 1834: 600 
(China). 

Lamprotula leaii (Gray in Griffith & Pidgeon, 
1833) — Haas, 1969: 282-283. 


Remarks 
Southeast Asia (Haas, 1969). Type material 
not found in 2006 in the BMNH. 


Unio tenuis 
Gray in Griffith & Pidgeon, 1833 


Unio tenuis Gray in Griffith & Pidgeon, 1833: 
pl. 24, fig. 2. 

Anodon tenuis Gray — Griffith & Pidgeon, 1834: 
595: 


Unio tenuis Gray — Griffith & Pidgeon, 1834: 
601, citing the same figure. 

Symphynota discoidea Lea, 1834: 75 [187], 
al: 151 A939: 

Cristaria (Pletholophus) discoidea discoidea 
(Lea, 1834) — Haas, 1969: 388-389. 


Remarks 

Listed as a synonym of Cristaria (Pletholophus) 
discoidea discoidea (Lea, 1834) by Haas (1969) 
from Southeast Asia. However, Gray’s species 
was published a year earlier; ICZN Code Art. 
23.9 might apply here. Symphynota discoidea 
Lea, 1834, is the type species of Pletholophus 
Simpson, 1900: 585, by original designation. 
Type material not found in 2006 in the BMNH. 


Mycetopodidae 


Anodon georginae 
Gray in Griffith & Pidgeon, 1833 


Anodon georginae Gray in Griffith & Pidgeon, 
1833: pl. 19, fig. 3 [legend]. 

A. georginae Gray - Griffith & Pidgeon, 1834: 
595 (Rivers of Paraguay). 

Anodontites (Anodontites) trigonus georginae 
(Gray in Griffith & Pidgeon, 1833) — Haas, 
1969: 561. 


Remarks 
Southern South America (Haas, 1969). Type 
material not found in 2006 in the BMNH. 


Anodon susannae 
Gray in Griffith & Pidgeon, 1833 


Anodon susannae Gray in Griffith & Pidgeon, 
1333: pl. 24) Ni № 

A. susannae Gray - Griffith & Pidgeon, 1834: 
595 (South America). 

Anodontites (Anodontites) exoticus susannae 
(Gray in Griffith & Pidgeon, 1833) — Haas, 
1969: 571-572. 


Remarks 

Coastal rivers of southern Brazil and Uru- 
guay (Haas, 1969). Type material not found in 
2006 in the BMNH. 


Hyriidae 


Unio childreni 
Gray in Griffith & Pidgeon, 1833 


Unio chilensis Gray, 1828: 7, “pl. 6, fig. 12” 
[pl. never issued; copy in the BMNH]. 


230 PETIT & COAN 


U. childreni Gray in Griffith & Pidgeon, 1833; 
pl. 20, fig. 1 [legend]. 

U. childreni Gray — Griffith & Pidgeon, 1834: 
600 (South America). 

Diplodon (Diplodon) chilensis (Gray, 1828) — 
Haas, 1969: 511-512. 


Remarks 

Synonym of Diplodon (Diplodon) chilensis 
(Gray, 1828) from Chile (Haas, 1969). On p. 
600 just after the name of this species appears 
“(unio chinensis)’; see also under this species 
name below. The holotype of Unio chilensis 
Gray, 1828, is BMNH 1986147; the “holotype” 
of its “var B” is BMNH 1986148. Type material 
of Unio childreni not found in 2006 in the BMNH. 


Unio chinensis 
Griffith & Pidgeon, 1834 


Unio chinensis Griffith & Pidgeon, 1834: 600 
[in synonymy]. 


Remarks 

This name is listed as a synonym of Unio 
childreni Gray in the index of figures in Griffith 
& Pidgeon, 1834, has never been subse- 
quently used, and was probably an error for 
chilensis. lt is therefore unavailable (ICZN 
Code Art. 11.6) and does not preoccupy Unio 
chinensis Lea, 1868: 150. 


Unio smithii 
Gray in Griffith & Pidgeon, 1833 


Unio chilensis Gray, 1828: 7, “pl. 6, fig. 12” 
[pl. never issued; copy in the BMNH]. 

U. smithii Gray in Griffith & Pidgeon, 1833: pl. 
20, fig. 3 [legend]. 

U. smithii Gray — Griffith & Pidgeon, 1834: 600. 

Diplodon (Diplodon) chilensis (Gray, 1828) — 
Haas, 1969: 511-512. 


Remarks 

Synonym of Diplodon (Diplodon) chilensis 
(Gray, 1828) (Haas, 1969) from southern 
Chile, including Isla Chiloe. Type material of 
Unio smithii not found in 2006 in the BMNH 


Crassatellidae 


Mesodesma ornata 
Gray in Griffith & Pidgeon, 1833 


Mesodesma ornata Gray in Griffith & Pidgeon, 
1833: pl. 22, fig. 6 [legend]. 
M. ornata Gray — Griffith & Pidgeon, 1834: 598. 


Crassatella ornata (Gray) — Reeve, 1842a: 46 
[repr.: 306]; 1843: pl. 3, fig. 17. 

С. ornata Reeve — Löbbecke & Kobelt, 1886: 
16, pl. 6, fig. 4. 


Remarks 

Eucrassatella ornata (Gray in Griffith & 
Pidgeon, 1833) from West Africa. Reeve 
(1842a, 1843) was not certain that the species 
he discussed and figured was the same as that 
of Gray. Löbbecke & Kobelt (1886) and Lamy 
(1917: 235-236) incorrectly attributed the spe- 
cies to Reeve (ICZN Code Art. 49), Lamy add- 
ing that it occurs in Mauritania and Senegal. 
Type material not found in 2006 in the BMNH. 


Glauconomidae 


Glycymeris apinensis 
Gray in Griffith & Pidgeon, 1833 


Glauconome chinensis Gray, 1828: 6—7, “pl. 3, 
fig. 13, 13a” [pl. never issued; copy in the ВММН]. 

Glycymeris apinensis Gray in Griffith & 
Pidgeon, 1833: pl. 31, fig. 2 [legend]. 

Glauconome chinensis (Gray, 1828) — Griffith 
& Pidgeon, 1834: 597, with a footnote 
“Named on the plate in error as Glycemeris 
[sic] apinensis.” 

С. chinensis Gray — Robba et al., 2002: 116- 
117, pl. 20, fig. 1a, b. 

G. chinensis Gray — Qi, 2004: 315. 


Remarks 

This is Glauconome chinensis Gray, 1828, 
from the western Pacific (Robba et al., 2002; 
Qi, 2004). It is possible that “apinensis” was а 
typographical error for “chinensis”. Holotype 
of G chinensis: BMNH 1981214. 


Pharidae 


Solen novaculina 
Gray in Griffith & Pidgeon, 1833 


Solen constrictus Lamarck, 1818: 455. 

S. novaculina Gray in Griffith & Pidgeon, 1833: 
pisa: shige sl. 

S. novaculina Gray — Griffith & Pidgeon, 1834: 
600. 


Remarks 

Probably a synonym of brackish-water Sino- 
novacula constrictus (Lamarck, 1818), from 
Southeast Asia, and having nothing to do with 
the related fresh-water genus Novaculina 
Benson, 1830: 63 [type species, by monotypy: 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 251 


Novaculina gangetica Benson, 1830], which is 
smaller and has a more oval shape. Type ma- 
terial not found in 2006 in the BMNH. 


Solen sayii 
Gray in Griffith & Pidgeon, 1833 


Solen costatus Say, 1822: 315. 
Non S. costatus (Schumacher, 1817: 42, 126). 
$. sayii Gray in Griffith & Pidgeon, 1833: pl. 
31, fig. 3 [legend]. 
S. sayii — Griffith & Pidgeon, 1834: 600. 


Remarks 

Not attributed to any author in the Index of 
Griffith & Pidgeon (1834), and not listed by 
Sherborn (1922-1932). Synonymized by 
Stimpson (1851: 22) with Siliqua costata (Say, 
1822). Type material not found in 2006 in the 
BMNH. It is possible that this name was meant 
as a replacement for Solen costatus Say, 1822, 
non S. costatus (Schumacher, 1817) [originally 
Leguminaria costatus]; the latter is now re- 
garded as a synonym of Solen radiatus 
Linnaeus, 1758: 673, the type species by 
monotypy of Siliqua Megerle von Mühlfeld, 
1811: 44. Presumably ICZN Code Art. 23.9.2 
would apply to conserve the name of Say’s 
well-known western Atlantic species. 


Solen tenuis 
Gray in Griffith & Pidgeon, 1833 


Solen tenuis Gray in Griffith & Pidgeon, 1833: 
pl. 31, fig. 4 [legend]. 

S. tenuis Gray — Griffith & Pidgeon, 1834: 600. 
Non Solen tenuis W. Wood, 1828b: 3, pl. 1 
Solen fig. 6; non Solen tenuis Broderip & G. 
B. Sowerby |, 1829: 361. 

S. africanus Chenu, 1843: Solen pl. 2, fig. 8, 8a. 

Sinupharus africanus (Chenu) — Cosel, 1993: 
247-250. 


Remarks 

Synonym of Sinupharus africanus (Chenu, 
1843) (Cosel, 1993). According to Cosel, the 
holotype of Gray’s species is in the BMNH, 
but it could not be found in the type or general 
collections in 2006. 


Mesodesmatidae 


Erycina denticulata 
Gray, 1825 


Mactra deaurata Turton, 1822: xxvi, 71. 
Erycina denticulata Gray, 1825: 135 [non 
Deshayes, 1856: 182]. 


Mactra denticulata — W. Wood, 1828b: 4, pl. 
1, Mactra fig. 9. 

Mesodesma denticulata — Griffith & Pidgeon, 
1682) pl. 22, fig. 2: 

M. denticulata (Gray) — Griffith 8 Pidgeon, 
1834: 598. 


Remarks 

This Gray species dates from 1825 and is a 
synonym of the northwestern Atlantic Meso- 
desma deauratum (Turton, 1822) (Lamy, 1914: 
19-20; Davis, 1965). Type material not found in 
2006 in the BMNH. 


Mactra subtriangulata 
W. Wood, 1828 


?Erycina subangulata Gray, 1825: 135, nom. 
nud., as possible synonym of Crassatella 
cuneata Lamarck, 1818: 483. 

Mactra subtriangulata W. Wood, 1828b: 4, pl. 
1, Mactra fig. 10. 

Mesodesma subtriangulata Gray - Griffith 8 
Pidgeon, 1833: pl. 22, fig. 1 [legend]. 

М. subtriangulata Gray — Griffith & Pidgeon, 
1834: 598. 

Paphies (Mesodesma) subtriangulata 
subtriangulata (W. Wood) — Powell, 1979: 
416, pl Se fig, 12. 

Paphies (Paphies) subtriangulatum (W. Wood, 
1828) — Beu & Rooij-Schuling, 1982: 214-218. 


Remarks 

Mesodesma subtriangulata was attributed to 
Gray in the list of figures in Griffith & Pidgeon 
(1834), but the name actually was made avail- 
able by Wood (1828b), who figured it from a 
BM specimen. The name may have come from 
Gray, who listed in synonymy three years ear- 
lier what may have been the same species 
name, but spelled “subangulata”. lt was also 
attributed to Gray by Hanley (1856: 202), who 
incorrectly listed it as a synonym of Crassatella 
cuneata Lamarck, 1818. This species occurs 
in New Zealand (Beu & Rooij-Schuling, 1982), 
who cited BMNH 19821 as “almost certain 
holotype”. This type lot contains two separate 
valves, the right valve figured in Griffith & 
Pidgeon; the other valve is a different species, 
according to Beu & Rooij-Schuling. There are 
three separate lots of uncertain status in the 
same box with these specimens. 


Erycina solenoides 
King & Broderip, 1832 


Erycina solenoides King & Broderip, 1832: 
335. 


232 PETIT & COAN 


Mesodesma solenoides Gray — Griffith & 
Pidgeon, 1834: 598. 

Darina solenoides (King & Broderip, 1832) — 
Forcelli, 2000: 162. 


Remarks 

Mesodesma solenoides was misattributed to 
Gray in the list of figures in Griffith & Pidgeon 
(1834). It is the Chilean Darina solenoides (King 
& Broderip, 1832) (Forcelli, 2000). Type spe- 
cies of Darina Gray, 1853: 42, by monotypy. 
The type material of Erycina solenoides is 
BMNH 1837.12.1.879 & 884, 2 syntypes; 
1859.9.19.59, 3 syntypes; BMNH 1968507, 2 
syntypes. 


Tellinidae 


Tellina guildfordiae 
Gray in Griffith & Pidgeon, 1833 


Tellina lutea W. Wood, 1828b: 3, pl. 1, Tellina 
fig. 3. 

T. guildfordiae Gray in Griffith & Pidgeon, 1833: 
pl. 19, fig. 2 [legend]. 

T. guildfordiae Gray — Griffith & Pidgeon, 1834: 
600. 

_Т. (Megangulus) lutea W. Wood, 1828b — Coan 

et al., 2000: 403. 


Remarks 

Synonym of Те/та (Megangulus) lutea W. 
Wood, 1828b (Coan et al., 2000). Type mate- 
rial not found in 2006 in the BMNH. 


Cuspidariidae 


Neaera chinensis 
Gray in Griffith & Pidgeon, 1833 


Neaera chinensis Gray in Griffith & Pidgeon, 
1833: pl. 22, fig. 5 [legend]. 

Neroea chinensis Gray — Griffith & Pidgeon, 
1834: 598. 

Cuspidaria chinensis (Griffith & Pidgeon) — 
Habe, 1977: 320. 

C. chinensis (Gray in Griffith & Pidgeon) — 
Poutiers & Bernard, 1995: 158. 

C. chinensis (Griffith & Pidgeon) — Wu, 2004: 
33: 


Remarks 

Cuspidaria chinensis (Gray in Griffith & 
Pidgeon, 1833) from the western Pacific (Habe, 
1977; Poutiers & Bernard, 1995; Wu, 2004), but 
not listed in Okutani (2000) or Xu (2004). Type 
species, by monotypy, of Neaera Gray in Griffith 


& Pidgeon, 1833, non Robineau-Desvoidy, 1830, 
a synonym of Cuspidaria Nardo, 1840: 202; see 
also under genus Neaera above. Cuspidaria 
corrugata Prashad, 1932 [pp. 329-330, pl. 7, fig. 
39], was originally compared to C. chinensis, 
Prashad noting that his species was less ros- 
trate, with less prominent umbones and less 
developed, less regular sculpture. Holotype of 
Gray’s species: BMNH 1996471. 


Gastropoda 
Trochidae 


Trochus bicarinatus 
Gray in Griffith & Pidgeon, 1833 


Trochus bicarinatus Gray in Griffith & Pidgeon, 
1833: pl. 1, fig. 1 [legend]. 

T. bicarinatus Gray — Griffith & Pidgeon, 1834: 
600. 
Non Trochus bicarinatus Lamarck, 1804: 50 
[and two other senior homonyms]. 

T. grayanus Philippi, 1850 [in 1846-1855]: pl. 
Adige; TODOS: 281, non nev pro f. 
bicarinatus Griffith & Pidgeon, non Lamarck. 


Remarks 

Listed as a synonym of the eastern Atlantic 
Gibbula magus (Linnaeus, 1758: 757) by 
Pilsbry (1889: 197; Sabelli et al., 1990: 131), 
the type species of Gibbula Risso, 1826: 134, 
by the subsequent designation of Herr- 
mannsen (1847: 473). However, this seems 
unlikely, because Gray’s taxon does not have 
the open umbilicus of this species of Gibbula. 
Type material not found in 2006 in the BMNH. 
It may best be regarded as a nomen dubium 
(J. H. McLean & В. А. Marshall, personal com- 
munications, 4 Jan. 2007). 


Calliostomatidae 


Trochus cunninghami 
Gray in Griffith & Pidgeon, 1833 


Trochus selectus Dillwyn, 1817: 801. 

T. cunninghami Gray in Griffith & Pidgeon, 
1833: pl. 1, fig. 7 [legend]. 

T. cunninghami Gray — Griffith & Pidgeon, 
1834: 600. 

Calliostoma (Maurea) selectum (Dillwyn, 
1817) — Marshall, 1995: 84, 108-110. 


Remarks 
Synonym of Calliostoma (Maurea) selectum 
(Dillwyn, 1817) (Marshall, 1995). Trochus 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 235 


cunninghami is the type species of СаШо- 
tropis Oliver, 1926: 110, by original designa- 
tion, which Marshall synonymized with 
Maurea Oliver, 1926: 107. Additionally, 
Calliotropis Oliver, 1926, is a junior homonym 
of Calliotropis Seguenza, 1903: 462, and was 
replaced by Calotropis Thiele, 1929: 49, 
which Marshall also placed into this generic 
synonymy. Holotype of Т. cunninghami: 
BMNH 1987047. 


Ampullariidae 


Paludina pulchra 
Gray in Griffith & Pidgeon, 1833 


Paludina pulchra Gray in Griffith & Pidgeon, 
1833: pl. 1, fig. 6 [legend]. 

P. pulchra Gray — Griffith & Pidgeon, 1834: 
599. 

Ampularia pulchra (Gray in Griffith & Pidgeon) 
— С. В. Sowerby Ш, 1909: 356. 

Pomacea (Pomacea) pulchra (Gray in Griffith 
& Pidgeon) — Cowie & Thiengo, 2003: 73. 


Remarks 

Pomacea (Pomacea) pulchra (Gray in Griffith 
& Pidgeon, 1833), from South America (Cowie 
& Thiengo, 2003). Holotype: BMNH 20020680. 


Cyclophoridae 


Cyclostoma auriculare 
Gray in Griffith & Pidgeon, 1833 


Cyclostoma auriculare Gray in Griffith & 
Pidgeon, 1833: pl. 28, fig. 5 [legend; first is- 
sue of pl. with reversed images]; 1834: pl. 
28*, fig. 5 [legend; second issue of pl. with 
corrected images]. 

C. auriculare Gray — Gray in Griffith & Pidgeon, 
1834: 587. 


Remarks 
Possibly an Otopoma. Type material not 
found in 2006 in the BMNH. 


Viviparidae 


Paludina chinensis 
Gray in Griffith & Pidgeon, 1833 


Paludina chinensis Gray in Griffith & Pidgeon, 
1833: ply 1, fie. a: 

P. chinensis Gray — Gray in Griffith & Pidgeon, 
1834: 599. 


Viviparus chinensis (Gray) — Yen, 1942: 190, 
200, реа 


Remarks 

Cipangopaludina chinensis (Gray in Griffith 
& Pidgeon, 1833) from southeast Asia and 
widely introduced elsewhere (Cowie, 1997: 
13). Type material not found in 2006 in the 
BMNH. Yen (1942) figured a specimen as 
“type”, one of three in a Reeve lot, but it is 
most unlikely that this specimen was seen by 
Gray. 


Paludina subcostata 
Gray in Griffith & Pidgeon, 1833 


Paludina subcostata Gray in Griffith & Pidgeon, 
1833: pl. 36, fig. 3 [legend; first issue of pl. 
with reversed images]; 1834: pl. 36*, fig. 3 
legend; second issue of pl. with corrected 
images]. 

Р subcostata Gray — Gray in Griffith & Pidgeon, 
1834: 596 (China). 

Viviparus subcostatus (Gray) — Yen, 1942: 192, 
201, pl. 14, fig. 36. 


Remarks 

Perhaps a Chinese Viviparus (Yen, 1942), 
who figured a specimen in the BMNH collec- 
tion as “holotype”. BMNH 20070171, probable 
syntype, ex [John] Reeves. 


Pachychilidae 


Melania carolinae 
Gray in Griffith & Pidgeon, 1833 


Melania costula Rafinesque, 1833: 166 
[Spring]. 

M. carolinae Gray in Griffith & Pidgeon, 1833 
[Dec.]: pl. 13, fig. 3 [legend]. 

M. carolinae Gray — Griffith & Pidgeon, 1834: 
598. 

Brotia costula (Rafinesque) — Köhler & 
Glaubrecht, 2001: 295-299, fig. 10. 

В. carolinae Gray — Köhler & Glaubrecht, 2002: 
130-131. 

В. costula (Rafinesque) — Kohler & Glaubrecht, 
2006: 176-181, figs. 17-19. 


Remarks 

Synonym of Brotia costula (Rafinesque, 
1833) from India and Southeast Asia (Köhler 
& Glaubrecht, 2001, 2006). Lectotype (Kohler 
& Glaubrecht, 2002) of M. carolinae: BMNH 
1874.10.12.11/1; /2, paralectotype. 


234 PETIT & COAN 


Melania freethii 
Gray, 1831 


Melania freethii Gray, 1831: 11. 

M. frethii Gray — Griffith & Pidgeon, 1833: pl. 
14, fig. 2 [legend]; 1834: 598. 

Potadoma freethii (Gray) — Morrison, 1954: 370. 

Р freethii (Gray) — Mandahl-Barth, 1967: 113-— 
ATS 

P. freethii (Gray) — D. S. Brown, 1994: 114. 


Remarks 

The entries in Griffith & Pidgeon are misspell- 
ings of Potadoma freethii (Gray, 1831) de- 
scribed from Fernando Ро [Bioko] Island, 
Equatorial Guinea, West Africa, and named 
for a Col. Freeth. It is the type species of 
Potadoma Swainson, 1840: 199, 341, by the 
subsequent designation of Gray (1847: 152), 
and occurs from the Ivory Coast to lower Zaire. 
BMNH 74.10.12.10, two specimens, the larger 
of which is labeled “holotype”. 


Melania henriettae 
Gray in Griffith & Pidgeon, 1833 


Melania henriettae Gray in Griffith & Pidgeon, 
1833: pl. 13, fig. 2 [legend]. 

M. henriettae Gray — Griffith & Pidgeon, 1834: 
598. 

Brotia henriettae Gray — Morrison, 1954: 383. 

В. henriettae Gray — Köhler & Glaubrecht, 
2001: 285, 309. 

В. henriettae Gray — Köhler & Glaubrecht, 
2002: 137-138. 


Remarks 

A valid Chinese species in genus Brotia 
(Köhler & Glaubrecht, 2001) and type species 
by original designation of Wanga Chen, 1943: 
20-21, a subjective synonym of Brotia H. 
Adams, 1866: 150. Yen (1942: 204, 216, pl. 
15, fig. 66) figured one of two specimens in a 
lot in the BMNH as “type”, and this specimen 
was designated as “lectotype” by Köhler & 
Glaubrecht, 2002: 137), BMNH 19990495/A, 
the other, /B, designated as “paralectotype”. 
According to Köhler & Glaubrecht (2002), this 
lot is labeled “China, leg. [John] Reeves”. 


Melania laevis 
Gray in Griffith & Pidgeon, 1833 


Melania laevis Gray in Griffith & Pidgeon, 
1833: pl. 14, fig. 8; 1834: 598. 


[Non Melania laevis Lea, 1843: 237, nom. 
nov. pro Melania laevigata Lea, 1841: 11, non 
Lamarck, 1822: 165]. 

M. laevis Gray — Griffith & Pidgeon, 1834: 598. 


Remarks 
Type material not found in 2006 in the 
BMNH. Possibly a Melanoides. 


Melania subcarinata 
Gray in Griffith & Pidgeon, 1833 


Melania subcarinata Gray in Griffith & Pidgeon, 
1833: pl. 14, fig. 7 [legend]. 

M. subcarinata Gray — Griffith & Pidgeon, 
1834: 598. 
[Non Melania subcarinata Reeve, 1860: pl. 
40, fig. 282, ex Anthony ms]. 


Remarks 

BMNH 74.10.12.8, 2 specimens, one of 
which is labeled “holotype”. Possibly a Pachy- 
chilus. 


Paludomidae 


Melania conica 
Gray in Griffith & Pidgeon, 1833 


Melania conica Gray in Griffith & Pidgeon, 
1833: pl. 14, fig. 5 [legend]. 
Non Melania conica Say, 1821: 176. 

M. conica Gray — Griffith & Pidgeon, 1834: 598 
(Ceylon). 


Remarks 

As Paludomus conica (Gray) in Reeve 
(1847b: pl. 3, fig. 14) from India. Gray's taxon 
is, however, a junior homonym, which has 
probably never been renamed, a task best 
undertaken by a reviser of this genus. Type 
material not found in 2006 in the BMNH. 


Melania globulosa 
Gray in Griffith & Pidgeon, 1833 


Melania globulosa Gray in Griffith & Pidgeon, 
1833: pl. 14, fig. 6 [legend]. 

M. globulosa Gray — Griffith & Pidgeon, 1834: 
589. 


Remarks 

As Paludomus globulosa (Gray) in Reeve 
(1847b: pl. 40, fig. 282) from Ceylon. Holo- 
type: BMNH 1912.1.26.1. 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 235 


Thiaridae 


Melania lineolata 
W. Wood, 1828 


Melania lineolata W. Wood, 1828b: 13, 42, pl. 
4, Strombus fig. 11; not pl. 7, Helix fig. 26, 
also incorrectly listed on p. 42 for this spe- 
cies, but then correctly shown on p. 22 as 
Helix laevissimus W. Wood, 1828b. 

M. lineolata - Griffith & Pidgeon, 1833: pl. 13, 
fig. 4. 

M. lineolata Gray — Griffith & Pidgeon, 1834: 
598. 

Melanopsis lineolatus (Gray) — Hanley, 1856, 
215, Suppl. pl. 4, Strombus fig. 11; listed as 
having been originally described as Melania. 

Aylacostoma (Hemisinus) lineolata (Gray) — 
Morrison, 1954: 376. 


Remarks 

This species was misattributed to Gray in 
the index to figures in Griffith & Pidgeon 
(1834), although it is possible that Gray is re- 
sponsible for the name first made available 
by Wood (1828). Wood’s taxon is the type 
species of Hemisinus Swainson, 1840: 199, 
341, by monotypy. Swainson (1840: 200), in 
describing this new genus, noted the type as 
Melania lineata* [sic], the asterisk referring to 
a footnote: “Gray, in Griff. Cuv. pl. 13, fig. 4.” 
Later (p. 341) he listed the species as “H. 
lineolata. Griff. Cuv. xii. pl. 13, f. 4”. This spe- 
cies occurs in Jamaica, and it is to be cited as 
Hemisinus lineolatus (W. Wood, 1828). 

Morrison (1954) misattributed this species 
to Gray, but dated it as 1828. Morrison’s ref- 
erences do not contain any work by Wood. 
However, he did list Gray, in Griffith & Pidgeon 
(1834). This was the only species he included 
that was in Griffith & Pidgeon. 

See below under Potamididae for a second 
Melania lineolata. 


Melania quadriseriata 
Gray in Griffith & Pidgeon, 1833 


Melania quadriseriata Gray in Griffith & 
Pidgeon, 1833: pl. 14, fig. 3. 

M. quadriseriata Gray — Griffith & Pidgeon, 
1834: 598. 


Remarks 
Type maierial not found in 2006 in the 
BMNH. 


Melania retusa 
Gray in Griffith & Pidgeon, 1833 


Melania retusa Gray in Griffith & Pidgeon, 
1833: pl. 14, fig. 9 [legend]. 
M. retusa Gray — Griffith & Pidgeon, 1834: 598. 


Remarks 
Type material not found in 2006 in the 
BMNH. 


Potamididae 


Melania lineolata 
Gray in Griffith & Pidgeon, 1833; 
Cerithium truncatum 
Griffith & Pidgeon, 1834 


Melania lineolata Gray in Griffith & Pidgeon, 
1833: pl. 14, fig. 4. 

Non Melania lineolata W. Wood, 1828b: 42, 
pl. 4, fig. 11; pl. 7, fig. 26 [see above under 
Thiaridae]. 

Cerithium truncatum Lam. [sic] — Griffith & 
Pidgeon, 1834: 596, ref. to pl. 14, fig. 4. 
Non Cerithium truncatum Gray in Griffith & 
Pidgeon, 1833: pl. 13, fig. 1. 

Cerethium [sic] truncatum Griffith & Pidgeon, 
1834: 598, reference for pl. 14, fig. 4, listed 
under Melania — “Melania lineolata, in plate, 
is Cerethium [sic] truncatum.” 

Non Cerithium truncatum Gray in Griffith & 
Pidgeon, 1833: pl. 13, fig. 1. 

Cerithidea reidi Houbrick, 1986: 280-286, figs. 

1-4, 9, 11-13. 


Remarks 

Renamed by Griffith & Pidgeon (1834), pre- 
sumably because of the earlier M. lineolata 
Gray [sic; =W. Wood] also figured in this work 
(see above under Thairidae). Perhaps Gray 
supplied Griffith & Pidgeon with the replace- 
ment name, but there is no explicit evidence 
for this. 

This is the type species of Cerithidea 
Swainson, 1840: 203, 342, by the subsequent 
designation of Makiyama, 1936: 331. Houbrick 
(1984: 15-16) and Wilson (1993: 132) thought 
this species was a synonym of Cerithidea 
obtusa (Lamarck, 1822: 71), but Lamarck’s 
species is broader anteriorly and has more 
prominent axial ribs (D. Reid, personal com- 
munication, December 2006). The subsequent 
designation by Pilsbry & Harbison (1933: 115) 
cited by some authors, such as Bequaert 


236 PETIT & COAN 


(1942: 20-21), is not valid because Pilsbry & 
Harbison did not mention either of the spe- 
cies originally included by Swainson (1840). 
Type material is not present in the BMNH. The 
name Cerithidea reidi proposed for this spe- 
cies is valid, because both Melania lineolata 
Gray in Griffith & Pidgeon, 1833, and the re- 
placement Cerithium truncatum Griffith & 
Pidgeon, 1834, are junior primary homonyms. 

Houbrick (1986) did not discuss the convo- 
luted history of the name Cerithium truncatum, 
but his description of the species as Cerithium 
reidi provides its only available name. The at- 
tribution of C. truncatum to Lamarck on p. 596 
must be in error; the species was never de- 
scribed by Lamarck. 


Campanilidae 


Cerithium truncatum 
Gray in Griffith & Pidgeon, 1833; 
C. laeve “Gray” 


Cerithium leve Quoy & Gaimard, 1834: 106- 
108; 1833: pl. 54, figs. 1-3 [pl. without leg- 
end]. 

Non Cerithium laevis Perry, 1810: pl. 15, fig. 
3 [ICZN Code Art. 58.1 — spellings differing 
only in e vs. ae are equivalent]. 

C. truncatum Gray in Griffith & Pidgeon, 1833: 
pl. 13, fig. 1 [legend]. 

C. laeve Gray — Griffith & Pidgeon, 1834: 596 

[with footnote stating that the plate had been 
miscaptioned as C. truncatum] (“New Hol- 
land”). 
Non Cerithium truncatum Lam. [sic] — Griffith 
& Pidgeon, 1834: 596 — Reference for pl. 
14, fig. 4 [= Melania lineolata Gray in Griffith 
& Pidgeon, 1833]. 

Campanile symbolicum lredale, 1917: 325- 
326, nom. nov. pro Cerithium leve Quoy & 
Gaimard, non Perry. 

С. symbolicum lredale — Houbrick, 1981: 282- 
283, 

C. symbolicum lredale — Houbrick, 1989: 1-6. 


Remarks 

Both Iredale (1917) and Houbrick (1981) re- 
jected the name truncatum for this species. It 
is not a junior homonym, and the fact that Gray 
decided that the name truncatum was un- 
needed because the species had already been 
named /eve by Quoy & Gaimard, not knowing 
that the latter was a junior homonym, does not 
make truncatum unavailable. However, as a 
result of this action by Iredale (1917), ICZN 


Code Art. 23.9 probably applies, and Iredale's 
unnecessary replacement name is now to be 
used. This unique Recent taxon occurs in 
southwestern Australia and is the type species, 
by monotypy, of Ceratoptilus Bouvier, 1887: 36, 
a junior synonym of Campanile Fischer, 1884: 
680. BMNH 2007013/1, probable figured 
syntype; BMNH 2007013/2, additional syntype; 
BMNH 20070102, additional Gray specimens 
of uncertain type status. 


Littorinidae 


Littorina pulchra 
С. В. Sowerby |, 1832 


Turbo zebra Donovan, 1825: pl. 130, [3] pp. 
expl. to “pl. 131”. 

Littorina pulchra G. B. Sowerby |, 1832a: 
Littorina, figs. 2, 3. 

Littoraria pulchra Gray — Griffith & Pidgeon, 
1834: 598. 

Littorina zebra (Donovan) — Keen, 1971: 366, 
369, fig. 189. 

Littoraria (Littoraria) zebra (Donovan) — Reid, 
1999: 35-38, figs. 2H, |, 3D, 4E, 6C, 7D—F, 
ЗЕ-Н. 


Remarks 

This eastern Pacific species was mis- 
attributed to Gray in the index to figures in 
Griffith & Pidgeon (1834). As shown, it was 
first named as “pulchra” by С. В. Sowerby 1, 
but was earlier described by Donovan, whose 
name has priority. It is the type species, by 
monotypy, of Littoraria Gray in Griffith & 
Pidgeon, 1833; see also under this genus 
above. The type material of Sowerby’s spe- 
cies is not present in the BMNH. 


Naticidae 


Natica bifasciata 
Gray in Griffith & Pidgeon, 1833 


Natica bifasciata Gray in Griffith & Pidgeon, 
1833: pl. 1. fig. 2. 

N. bifasciata Gray — Griffith 4 Pidgeon, 1834: 
598. 

Polinices (Polinices) bifasciata Griffith & Pidgeon, 
1833 — Keen, 1971: 476, 478, fig. 873. 


Remarks 

This species occurs in the Panamic prov- 
ince (Keen, 1971). Type material not present 
in the BMNH. 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 237 


Cryptosoma javanica 
Gray in Griffith & Pidgeon, 1834 


Cryptosoma javanica Gray in Griffith & 
Pidgeon, 1834: pl. 41, fig. 1 [legend]; 596. 
C. javanica Gray — Griffith & Pidgeon, 1834: 
596. 

Sinum javanicum Gray in Griffith & Pidgeon — 
Swennen et al., 2001: 121. 

S. javanicum Griffith & Pidgeon — Ma, 2004: 65. 


Remarks 

Sinum javanicum (Gray in Griffith & Pidgeon, 
1834) from Southeast Asia. Type material not 
present in the BMNH. 


Pomatiasidae 


Cyclostoma articulata 
Gray in Griffith & Pidgeon, 1833 


Cyclostoma articulata Gray in Griffith & 
Pidgeon, 1833: pl. 28, fig. 1 [legend; first is- 
sue of pl. with reversed images]; 1834: pl. 
28*, fig. 1 [legend; second issue of pl. with 
corrected images]. 

С. articulata Gray — Griffith & Pidgeon, 1834: 
596. 


Remarks 

Tropidophora articulata (Gray in Griffith & 
Pidgeon, 1833) from Rodrigues and Mauritius 
islands (Abbott, 1989: 48). BMNH 20070165, 
probable syntype, found during this review. 


Cyclostoma pulchrum 
Gray in Griffith & Pidgeon, 1833 


Cyclostoma pulchra Gray in Griffith & Pidgeon, 
1833: pl. 28, fig. 3 [legend; first issue of pl. 
with reversed images]; 1834: pl. 28*, fig. 3 [leg- 
end; second issue of pl. with corrected images]. 

C. pulchra Gray — Griffith & Pidgeon, 1834: 
596 [as С. pulchrum)]. 

Non Cyclostoma pulchrum W. Wood, 1828b: 
36, pl. 6, Turbo fig. 4. 

Tropidophora pulchra (Gray) — Gerlach, 2006: 

29.1258] 261] 


Remarks 

Because the Greek noun —stoma, mouth, is 
neuter, the species name as originally pro- 
posed should have been pulchrum, and it was 
so corrected in the Index on p. 596. BMNH 
20070169, is of doubtful type status. This spe- 
cies occurs on the Seychelles (Gerlach, 2006). 


Three separate taxa are involved here in a 
nomenclatural tangle: (1) Turbo pulcher Dillwyn, 
1817, a marine snail, in the family Potamididae; 
(2) Turbo pulcher W. Wood, 1828b, a Jamai- 
can land snail; (3) Cyclostoma pulchrum Gray 
in Griffith & Pidgeon, 1833, the Seychelles land 
snail. 

The morphologically similar Jamaican spe- 
cies Megannularia pulchra (W. Wood, 1828b) 
has sometimes been confused with Gray’s 
taxon (e.g., Reeve, 1842b: 98). Wood intro- 
duced the new name Turbo pulcher Wood (W. 
Wood, 1828b: 18, pl. 6, fig. 4) for the Jamaican 
species and later in the same work (p. 36) indi- 
cated that the applicable Lamarckian genus for 
the species was Cyclostoma. The confusion 
was later compounded by Sherborn, who mis- 
takenly listed Cyclostoma pulchrum (Dillwyn, 
1817: 855), a species of Potamididae originally 
described as Turbo pulcher Dillwyn, as having 


been transferred by Wood to Cyclostoma. 


Sherborn overlooked the fact that Wood intro- 
duced Turbo pulcher as a new species although 
he had also earlier listed and figured the dis- 
similar Turbo pulcher Dillwyn (Wood, 1828a: 
149, pl. 31, fig. 93). 

Homonymy does not exist between the 
Wood and Gray names as they were not origi- 
nally described in the same genus and are not 
now placed in the same genus. 

Wood’s somewhat confusing manner of list- 
ing names is discussed in Appendix B herein. 


Cyclostoma madagascariensis 
Gray in Griffith & Pidgeon, 1833 


Cyclostoma madagascariensis Gray in Griffith 
& Pidgeon, 1833: pl. 28, fig. 4 [legend; first 
issue of pl. with reversed images]; 1834: pl. 
28”, fig. 4 [legend; second issue of pl. with 
corrected images]. 

C. madagasciensis [sic] Gray — Griffith & 
Pidgeon, 1834: 597. 


Remarks 

Tropidophora madagascariensis (Gray in 
Griffith & Pidgeon). BMNH 20070170, 2 speci- 
mens of doubtful type status. 


Strombidae 


Strombus campbelli 
Gray in Griffith & Pidgeon, 1833 


Strombus campbelli Gray in Griffith & Pidgeon, 
1833: pl. 25, fig. 6 [legend]. 


238 PETIT & COAN 


S. campbelli Gray — Griffith & Pidgeon, 1834: 
600. 

S. (Doxander) vittatus campbelli Griffith & 
Pidgeon — Abbott, 1960: 111, 114-115. 


Remarks 

Strombus (Doxander) vittatus campbelli Gray 
in Griffith & Pidgeon, 1833, from northern Aus- 
tralia (Abbott, 1960). Type material not present 
in the BMNH. 


Strombus deformis 
Gray in Griffith & Pidgeon, 1833 


Lambis plicata Röding, 1798: 65. 

Strombus deformis Gray in Griffith & Pidgeon, 
1833: pl. 25, fig. 5 [legend]. 

S. deformis Gray — Griffith & Pidgeon, 1834: 
600. 

S. (Dolomena) plicatus plicatus (Réding, 1798) 
— Abbott, 1960: 89-90. 


Remarks 

Synonym of Strombus (Dolomena) plicatus 
plicatus (Rôding, 1798) (Abbott, 1960) from 
the Red Sea. BMNH 1953.3.11.39, holotype 
of S. deformis. 


?Epitoniidae 


Turritella sulurnalis 
Gray in Griffith & Pidgeon, 1833 


Turritella sulurnalis Gray in Griffith & Pidgeon, 
1833.21: ТЭН 

T. suturnalis “Sav.” — Griffith & Pidgeon, 1834: 
600. 

Eglisia tricarinata A. Adams & Reeve in Reeve, 
1849: pl. 1, fig. 3. 


Remarks 

Spelling changed and attributed in error to 
“Sav.” [Savigny] in the index of illustrations. This 
might well be in error for “Sow” [Sowerby], but 
it would have been a manuscript name, be- 
cause G. B. Sowerby | did not describe the spe- 
cies. Neither name is in Sherborn (1922-1932). 
Possibly an earlier name for Eglisia tricarinata 
A. Adams & Reeve in Reeve, 1849. The latter 
has generally been cited as 1850 from the Voy- 
age of the Samarang, but the species first ap- 
peared in 1849 in the Conchologia Iconica. The 
figure therein is not as good as that in the 
Samarang (A. Adams & Reeve, 1850: 49, pl. 
12, fig. 8). Type material of Gray’s species not 


found in 2006 in the BMNH housed as either 
Turritellidae or Epitoniidae. 


Buccinidae 


Triton elegans 
Gray in Griffith & Pidgeon, 1833 


Triton (Pusio) elegans Gray in Griffith & 
Pidgeon, 1833: pl. 25, fig. 2 [legend]. 

T. (Pusio) elegans Gray - Griffith & Pidgeon, 
1834: 600. 

Cantharus (Gemophos) elegans (Griffith & 
Pidgeon, ex Gray ms) — Keen, 1971: 559- 
2009-1108: 

Pusio elegans (Gray in Griffith & Pidgeon) — 
Vermeij, 2006: 86-87. 


Remarks 

This is the Panamic Pusio elegans (Gray in 
Griffith & Pidgeon, 1833) (Vermeij, 2006). See 
also Remarks under the genus Pusio above. 
BMNH 20070164, two possible syntypes. 


Triton iostoma 
Gray in Griffith & Pidgeon, 1833 


Triton iostoma Gray in Griffith & Pidgeon, 1833: 
pl. 23, fig. 4 [legend]. 

T. озюта Gray — Griffith & Pidgeon, 1834: 600. 

Pollia iostoma (Gray) — Gray, 1839: 112. 

Phos billenheusti Petit de la Saussaye, 1853: 
244, pl. 8, fig. 5. 

Cantharus (Prodotia) iostoma (Gray in Griffth 
& Pidgeon, 1834) — Cernohorsky, 1975: 200- 
203; 1986: 59-61. 


Remarks 

Cernohorsky (1986: 61) noted that the type 
is lost and designated Griffith & Pidgeon’s fig- 
ure as the “illustrated lectotype”. Type mate- 
rial not found in 2006 in the BMNH. A senior 
subjective synonym of Phos billeheusti Petit 
de la Saussaye, 1853, which is the type spe- 
cies, by original designation, of Prodotia Dall, 
1924: 89. There are several additional syn- 
onyms (Cernohorsky, 1975, 1986). Vermeij 
(2006: 86) regarded Prodotia as a full genus. 
Pollia Gray in С. В. Sowerby I, 1834: pl. 237, 
in which Gray placed this species in 1839, is 
now regarded as a different genus; its type 
species, by monotypy, is Buccinum undosum 
Linnaeus, 1758: 740 (Vermeij, 2006: 85-86). 
Prodotia iostoma occurs from East Africa to 
French Polynesia and the Hawaiian Islands. 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 239 


Triton nassoides 
Gray in Griffith & Pidgeon, 1834 


Triton nassoides Gray in Griffith & Pidgeon, 
1834: pl. 41, fig. 4. 

T. nassoides Gray — Griffith & Pidgeon, 1834: 
600. 


Remarks 

This is Nassaria nassoides (Gray in Griffith 
& Pidgeon, 1834) from the Philippine Islands. 
This species was overlooked by Sherborn 
(1922-1932). Type material not found in 2006 
in the BMNH. 

Triton nassoides Gray in Griffith & Pidgeon, 
1834, was one of 15 nominal species placed 
in the synonymy of Nassaria acuminata 
(Reeve, 1844) by Cernohorsky (1981: 17-21). 
In utilizing the later name for the species, he 
stated: “The type of Triton nassoides Griffith 
& Pidgeon, is lost, but the illustration shows 
that it is conspecific with Nassaria acuminata 
(Reeve). Since the taxon has not been used 
even once for a valid species since the time 
of description, it qualifies as a nomen oblitum.” 

There are two significant problems with 
Cernohorsky’s statement. First, in 1981 it was 
not possible for a name to be relegated to 
nomen oblitum status unilaterally, and action 
by the Commission was required. (There was 
a short period, from December 1970 to De- 
cember 1972, when such action had been 
possible. For comments and references about 
this misuse and misunderstanding of the ICZN 
Code, see Petit, 1987: 340). Cernohorsky was 
not alone in the misuse of nomina oblita. The 
provisions for nomina oblita have been 
changed again in the current Code (ICZN, 
1999), but Triton nassoides Gray in Griffith & 
Pidgeon does not qualify for “reversal of pre- 
cedence”, because it has been used as a valid 
name after 1899 (ICZN Code Article 23.9.1.1). 

Additionally, Cernohorsky’s statement that 
“the taxon has not been used even once for a 
valid species since the time of description” is 
inaccurate. There are at least eight usages, 
three of which not only figure Triton nassoides 
Gray in Griffith & Pidgeon but are in references 
that Cernohorsky listed in his chresonymy of 
Nassaria acuminata (Reeve), including Reeve 
(1844: Triton pl. 20, fig. 96 [correctly attributed 
to Gray, with reference to Griffith's Cuvier’s 
Animal Kingdom, in the text but to Reeve in 
the index]); С. В. Sowerby И (1859: 85, pl. 220, 
fig. 4 [misattributed to Reeve]); and Tryon 
(1881: 222, pl. 84, figs. 549, 550). Two post- 
1899 citations have also been located, both in 


lists of Philippine taxa. One of these, Hidalgo 
(1904: 43), lists as Hindsia nassoides Gray 
with a reference to the figure in G. B. Sowerby 
|| (1859). Even though he cited Sowerby, who 
incorrectly attributed the name to Reeve, 
Hidalgo correctly gave Gray as the author. 
Hidalgo listed four species of Hindsia in bold 
face and also listed, in normal type, “otras 
especies citadas.” These other species include 
Hindsia acuminata Reeve. The most recent 
citation located thus far is Faustino (1928: 
254), who listed the species as Nassaria 
nassoides Gray and included references to the 
three aforementioned works in which the spe- 
cies was figured. Faustino did not list Nassaria 
acuminata (Reeve), but included that name 
under Nassaria bitubercularis (A. Adams, 
1851: pl. 10, fig. 6, as Hindsia), with a refer- 
ence to Tnyon'(1881: 221). 


Nassa northiae 
Gray in Griffith & Pidgeon, 1833 


Nassa northiae Gray in Griffith & Pidgeon, 
1833: pl. 30, fig. 2 [legend]. 

N. northiae Gray - Griffith & Pidgeon, 1834: 
598. 

Northia northiae (Griffith & Pidgeon) — Keen, 
1971: 567-568, fig. 1136. 


Remarks 

This is a Panamic species (Keen, 1971). 
Type material not located in the BMNH in 2006. 
Oddly, in proposing the genus Northia, Gray 
(1847: 140) chose a different Panamic spe- 
cies, Buccinum pristis Deshayes, 1844: 192, 
as its type species by monotypy. 


Triton turbinelloides 
Gray in Griffith & Pidgeon, 1833 


Murex vibex Broderip, 1833a: 175 [14 Jan.]. 

Triton turbinelloides Gray in Griffith & Pidgeon, 
1833 [Dec.]: pl. 25, fig. 1 [legend; as subge- 
nus Pusio]. 

T. turbinelloides Gray — Griffith & Pidgeon, 
1834: 600. 

Cantharus (Gemophos) vibex (Broderip) — 
Keen, 1971: 561—562, fig. 1116. 

Hesperisternia vibex (Broderip, 1833) — 
Vermeij, 2006: 81. 


Remarks 

Synonym of the Panamic Hesperisternia 
vibex (Broderip, 1833) (Keen, 1971; Vermeij, 
2006). Holotype BMNH 20070168, possible 


syntype. 


240 PETIT & COAN 


Triton vexillum 
Gray in Griffith & Pidgeon, 1833 


Triton (Pusio) vexillum Gray in Griffith & 
Pidgeon, 1833: pl. 37, fig. 5 [legend]. 

T. vexillum Gray — Griffith & Pidgeon, 1834: 
600. 

Buccinum fasciculatum Reeve, 1846: pl. 10, 
fig. 76. 

Pisania crenilabrum A. Adams, 1855: 138. 

P. montrouzieri Crosse, 1862: 251, pl. 10, fig. 
5; 

P. crenilabrum A. Adams — Cernohorsky, 1971: 
138, figs. 3-9. 

P. fasciculatum Reeve — Cernohorsky, 1971: 
140, fig. 10. 

P. fasciculatum (Reeve) — Wilson, 1994: 96, 
pr: 10 HO 22am: 


Remarks 

The last three synonyms listed above were 
placed in synonymy by Wilson (1994). Earlier, 
Cernohorsky (1971) treated Pisania creni- 
labrum A. Adams as a valid species, with P 
montrouzieri Crosse in synonymy. At the same 
time, Cernohorsky (1971) treated Pisania 
fasciculata (Reeve) as a valid taxon and in its 
chresonymy listed “? Triton (Pusio) vexillum 
Gray in Griffith & Pidgeon”, commenting that 
the latter “is an undetermined species which 
closely resembles beach-worn specimens of 
Pisania fasciculata’. Species of Pisania are 
quite variable, but the specimen figured in color 
by Wilson does not appear to be separable 
from Gray’s Triton vexilium, which is thus prob- 
ably the senior synonym. The species occurs 
in the southwestern Pacific. Type material not 
found in 2006 in the BMNH. 

The last synonym listed, Pisania mon- 
trouzieri Crosse, 1862, is the type species, by 
monotypy, of Appisania Thiele, 1929: 314 (see 
also Boss & Bieler, 1991: 15), which Cerno- 
horsky (1971: 140) regarded as a synonym of 
Pisania Bivona-Bernardi, 1832: 8. 


Columbellidae 


Columbella suturalis 
Gray in Griffith & Pidgeon, 1834 


Columbella fluctuata G. B. Sowerby |, 1832b: 
115. 

C. suturalis Gray in Griffith & Pidgeon, 1834: 
pl. 41, fig. 3 [legend]. 

С. suturalis Gray — Griffith & Pidgeon, 1834: 
596. 


Anachis ( Costoanachis) fluctuata (G. B. Sower- 
by |, 1832b)-Keen, 1971: 578-579, fig. 1178. 


Remarks 

Synonym of the Panamic Anachis (Costo- 
anachis) fluctuata (С. В. Sowerby |, 1832b) 
(Keen, 1971). Type material of Gray’s species 
not found in 2006 in the BMNH. 


Columbella tylerae 
Gray in Griffith & Pidgeon, 1833 


Columbella tylerae Gray in Griffith & Pidgeon, 
1833: pl. 37, fig. 1 [legend; first issue of pl. 
with reversed images]; pl. 37%, fig. 1 [leg- 
end; second issue of pl. with corrected im- 
ages]. 

C. tylerae Gray — Griffith & Pidgeon, 1834: 596. 

Pyrene testudinaria tyleria [sic] Griffith & 
Pidgeon — Hu & Tao, 1995: 115, pl. 55, figs. 
8-14. 


Remarks 
This species has been reported from Taiwan. 
Type material not found in 2006 in the BMNH. 


Fasciolariidae 


Turbinella tubercularis 
Gray in Griffith & Pidgeon, 1833 


Turbinella tuberculata Broderip, 1833b: 7 [May 
17]. 

T. tubercularis — Gray in Griffith & Pidgeon, 
1833 [Dec.]: pl. 30, fig. 3. 

T. tubercularis Sav. [sic] — Griffith & Pidgeon, 
1834: 600. 

Leucozonia tuberculata (Broderip, 1833) — 
Keen, 1971: 615-616, fig. 1338. 


Remarks 

Attributed in error to “Sav.” [Savigny] in the 
list of illustrations. This was evidently in error 
for “Sow.” [G. В. Sowerby ||, and the species 
name a spelling error for the Panamic 
Turbinella tuberculata Broderip, 1833. The 
type material not found in 2006 in the BMNH. 


Nassariidae 


Cyllene owenii 
Gray in Griffith & Pidgeon, 1834 


Cyllene owenii Gray in Griffith & Pidgeon, 
1834: pl. 41, fig. 2 [legend]. 
C. owenii Gray — Griffith & Pidgeon, 1834: 597. 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 241 


С. owenii Gray in Griffith & Pidgeon — 
Cernohorsky, 1975: 166-167, figs. 90, 91 
[possible type figured]. 

С. owenii Gray — Kaicher, 1985: 4151 [syntype 
figured]. 


Remarks 

This is the type species, by monotypy, of 
Cyllene Gray in Griffith & Pidgeon, 1834; see 
under this genus above. This nassariid occurs 
in West Africa (Cernohorsky, 1975). BMNH 
1985006, 16 syntypes; label adds “Capt. E. 
Owen, R.N.” 


Bullia semiplicata 
Gray in Griffith & Pidgeon, 1833 


Bullia semiplicata Gray in Griffith & Pidgeon, 
1833: pl. 37, fig. 8 [legend; first issue of pl. 
with reversed images]; 1834: pl. 37*, fig. 8 [leg- 
end; second issue of pl. with corrected images]. 

Bullaea semiplicata Gray — Griffith & Pidgeon, 
1834: 596. 

Bullia semiplicata Gray — Gray, 1839: 127. 

В. semiplicata Gray, 1839 — Bosch € Bosch, 
1982: 103 [figured]. 


Remarks 

Type species, by monotypy, of Bullia Gray in 
Griffith & Pidgeon, a valid genus of the Nassarii- 
dae; see also under this genus above. Gray 
described, but did not figure, the species in 1839, 
with no indication that it was either new or had 
previously been described. This species occurs 
in the Gulf of Oman. BMNH 20070167, prob- 
able syntype; other material stored in the type 
collection was from Cuming and are not types. 


Melongenidae 


Fusus Striatus 
G. B. Sowerby |, 1833 


Fusus striatus С. В. Sowerby |, 1833: Fusus 
fig. 4. 
Non Fusus Striatus Réding, 1798: 119. 

Pyrula striata — Griffith & Pidgeon, 1833: pl. 
37, fig. 4 [legend; first issue of pl. with re- 
versed images]; 1834: pl. 37”, fig. 4 [legend; 
second issue of pl. with corrected images]. 

P. striata Sav. [sic] — Griffith & Pidgeon, 1834: 
599. 

P. clavella Reeve, 1847a: pl. 3, fig. 10. 

Taphon striatum (С. В. Sowerby |, 1833) — 
Bosch et al., 1995: 136, fig. 575. 


Remarks 

Pyrula striata is attributed on p. 599 to “Sav.” 
[Savigny], an error for “Sow.” [Sowerby]. Type 
species, by monotypy, of Taphon H. Adams & 
A. Adams, 1853: 151, who credited the spe- 
cies to Gray. Figured by Bosch et al. (1995) 
as Taphon striatum (G. B. Sowerby |, 1833), 
from the Red Sea. Unfortunately, Sowerby’s 
name is a junior primary homonym that has 
not been widely used, and the next available 
name, Pyrula clavella Reeve, 1847, should be 
used. 

Reeve's action in naming Pyrula clavella is 
inexplicable. In his discussion he stated: “An 
interesting species not included in M. Kiener’s 
monograph nor hitherto described; it is figured 
in Griffith's Cuvier PI. 37. Fig. 4. under the 
name of Р striata from which, it is scarcely 
necessary to add, it is quite distinct.” 

Exactly what P striata from which it is “quite 
distinct” is not stated, nor does Reeve explain 
the differences. Reeve did not treat any spe- 
cies as Pyrula striata. He also did not mention 
or figure Fusus striata Sowerby, which can- 
not, by any stretch of the imagination, be said 
to differ, because Sowerby’s figure is identical 
to that of Reeve. Reeve could not have had 
Fusus Striatus Réding in mind, because he did 
not know Réding’s publication existed. 


Muricidae 


Pyrula mawae 
Gray in Griffith & Pidgeon, 1833 


Pyrula mawae Gray in Griffith & Pidgeon, 
18337 pl. 25; figs.3, 4: 

P. mawae Gray - Griffith & Pidgeon, 1834: 599. 

Р. maweae [sic] Gray — Gray, 1839: 115 
(China). 

Latiaxis mawae (Griffith & Pidgeon) — 
Tsuchiya, in Okutani, 2000: 404—405. 


Remarks 

Latiaxis mawae (Gray in Griffith & Pidgeon) 
is a well-known Japanese species (Tsuchiya, 
in Okutani, ed., 2000). Gray’s later (1839) 
treatment suggests that his intention was to 
name the species for John Mawe’s wife, who 
contributed material to the BMNH and whose 
extensive collection was utilized by W. Wood. 
Unfortunately, the ICZN Code Art. 32 does not 
permit such orthographic errors to be cor- 
rected. Type material not found in 2006 in the 
BMNH. 


242 PETIT & COAN 


Mitridae 


Mitra chinensis 
Gray in Griffith & Pidgeon, 1834 


Mitra chinensis Gray in Griffith & Pidgeon, 
1834: pl. 40, fig. 2 [legend]. 

M. chinensis Gray — Griffith & Pidgeon, 1834: 
598. 

M. chinensis Gray — Gray, 1839: 135, pl. 35, 
ПО 

М. chinensis Gray — Yen, 1942: 236, 248, pl. 
24, fig. 172. 

M. chinensis Griffith & Pidgeon — Cernohorsky, 
1976: 331-332, pl. 284, figs. 1-6. 

M. chinensis Griffith & Pidgeon — Turner, 1993: 
83, 86-87. 

M. chinensis Gray — Li, 2004: 102, pl. 61C. 


Remarks 

Mitra chinensis Gray in Griffith & Pidgeon, 
1834 (Li, 2004, who attributed the species to 
Gray, 1839), from China. Sherborn (1930: 
5981) also attributed this species to Gray 
(1839). BMNH 1967.708/1-2, lectotype 
(Cernohorsky, 1976) and paralectotype. 


Mitra orientalis 
Gray in Griffith & Pidgeon, 1834 


Mitra orientalis Gray in Griffith & Pidgeon, 
1834: pl. 40, fig. 5 [legend]. 

M. orientalis Gray — Griffith & Pidgeon, 1834: 
598. 

M. maura Broderip, 1836: 193. 

M. orientalis Gray in Griffith & Pidgeon, 1834 
— Cernohorsky, 1970: 35. 

M. orientalis Griffith & Pidgeon — Cernohorsky, 
1976: 361-362, pl. 256, fig. 1, pl. 315, figs. 
1-6. 


Remarks 

Mitra orientalis Gray in Griffith & Pidgeon, 
1834, from northwestern South America 
(Cernohorsky, 1970). BMNH 1966417, -418, - 
419, syntypes, which are also syntypes of 
Mitra maura Broderip. There are additional 
junior synonyms (Cernohorsky, 1976). 


Volutidae 


Voluta broderipii 
Gray in Griffith & Pidgeon, 1833 


Voluta broderippii Gray in Griffith & Pidgeon, 
1833: pl. 26 [legend]. 


V. broderipi Gray — Griffith & Pidgeon, 1834: 
601. 

Melo (Melocorona) broderipii (Gray in Griffith 
& Pidgeon) — Weaver & duPont, 1970: 73, 
pl. 29, figs. C, D. 


Remarks 

This species occurs in the IndoPacific and 
is type species, by original designation, of 
Melocorona Pilsbry & Olsson, 1954: 24—25. 
Named for William Broderip, the incorrect 
double-p spelling on the 1833 plate was cor- 
rected in the 1834 list of figures. The spelling 
broderipii can be retained under ICZN Code 
Article 33.3.1, because it is an incorrect sub- 
sequent spelling in prevailing usage and is 
therefore deemed to be the correct original 
spelling. BMNH 1837,12,1.259, in the type 
collection, from Broderip, does not match the 
figure and is probably not the type (J. 
Pickering, personal communication, May 
2007). 


Voluta georginae 
Gray in Griffith & Pidgeon, 1833 


Voluta amphora [Lightfoot], 1786: 30. 

V. georginae Gray in Griffith & Pidgeon, 1834: 
pl. 34 [legend]. 

V. georginae Gray — Griffith & Pidgeon, 1834: 
601 (Swan River). 

Melo amphora ([Lightfoot], 1786) — Wilson, 
1994, 2: 125, pl. 26, figs. 1-6; pl. 26A: fig. 3; 
planas: 


Remarks 

Synonym of Melo amphora ([Lightfoot], 
1786) from Western and northern Australia 
(Wilson, 1994). BMNH 1952.8.1/1, holotype 
of V. georginae. 


Voluta miltonis 
Gray in Griffith 8 Pidgeon, 1833 


Voluta miltonis Gray in Griffith & Pidgeon, 
1833: pl. 29 [legend]. 

V. miltonis Gray — Griffith & Pidgeon, 1834: 
597 (New Holland). 

Melo miltonis (Gray in Griffith 8 Pidgeon) — 
Wilson, 1994: 125, pl. 27, figs. 2, 4, 6, 7; pl. 
26A, fig. 2. 


Remarks 

This species occurs in South and Western 
Australia (Wilson, 1994). BMNH 1952.5.12/2, 
holotype. 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 243 


Voluta pallida 
Gray in Griffith & Pidgeon, 1833 


Voluta pallida Gray in Griffith & Pidgeon, 1833: 
pl. 30, fig. 4. 

V. pallida Gray — Griffith & Pidgeon, 1834: 601 
(with “Vol. Grayii, Sow.” in synonymy). 
Non Voluta pallida Linnaeus, 1758: 727, type 
species, by monotypy, of the marginellid ge- 
nus Hyalina Schumacher, 1817 (Coan & Roth, 
1976). 

Атопа (Amoria) grayi Ludbrook, 1953: 136- 
137, pl. 14, figs. 4, 5, nom. nov. pro V. pallida 
Gray in Griffith & Pidgeon, non Linnaeus. 


Remarks 

Amoria (Amoria) grayi Ludbrook occurs in 
Western Australia (Ludbrook, 1953). Ludbrook 
chose the Sowerby manuscript name from the 
Index of Griffith & Pidgeon (1834) as the re- 
placement name. BMNH 1952.3.21/1, holotype. 


Voluta rudis 
Gray in Griffith & Pidgeon, 1833 


Voluta ferussacii Donovan, 1824: pl. 67, [3] 
рр. expl. 

V. rudis Gray in Griffith & Pidgeon, 1833: pl. 
30:11 1. 

V. rudis Gray — Gray in Griffith & Pidgeon, 
1834: 601. 

Adelomelon ferussacii (Donovan) — Weaver & 
duPont, 1974: 108; pl. 45, figs. C, D. 


Remarks 

Synonym of the Magellanic Adelomelon 
ferussacii (Donovan, 1824) (Weaver & duPont, 
1974). BMNH 1992177, holotype of V. rudis. 


Voluta turneri 
Gray in Griffith & Pidgeon, 1834 


Voluta turner Gray in Griffith 8 Pidgeon, 1834: 
pl. 40, fig. 1 [legend]. 

V. turneri Gray — Griffith & Pidgeon, 1834: 601. 

Amoria turneri (Gray in Griffith & Pidgeon) — 
Wilson, 1994, 2: 113, pl. 19, fig. 6a, b. 


Remarks 
This species occurs in northern Australia 
(Wilson, 1994). BMNH 1952.3.21/4, holotype. 
Olividae 


Ancillaria australis 
G. B. Sowerby |, 1830 


Ancillaria australis G. B. Sowerby |, 1830: 7, 
figs. 44—46. 

Ancillaria [no species name given], 1833: pl. 
37, fig. 6 [legend; first issue of pl. with re- 
versed images]; 1834: pl. 37*, fig. 8 [legend; 
second issue of pl. with corrected images, 
also without species name]. 

A. australis Sav. — Griffith & Pidgeon, 1834: 
595 [citing pl. 37, fig. 6]. 

Amalda australis (С. В. Sowerby) — Powell, 
1979: 208, pl. 43, fig. 17. 


Remarks 

Attribution of this name by Griffith & Pidgeon 
to “Sav.” [Savigny], was probably an error for 
“Sow.” [Sowerby]. Judging by illustrations, 
what was figured by Griffith & Pidgeon (1833) 
is a different species than Ancillaria australis 
С. В. Sowerby I, 1830, which actually occurs 
in New Zealand. This is best treated as a 
misidentification, rather than as a new spe-. 
cies and therefore a junior homonym. 


Terebridae 


Terebra africana 
Gray in Griffith & Pidgeon, 1833 


Terebra africana Gray in Griffith & Pidgeon, 
1833 [Dec.]: pl. 23, fig. 5 [legend]. 

T. africana Gray — Griffith & Pidgeon, 1834: 
600. 

T. variegata Gray, 1834: 61 [25 Nov.], nomen 
conservandum. 

T. variegata Gray — Keen, 1971: 685, 686, fig. 
1871, 

T. variegata Gray — Bratcher 8 Cernohorsky, 
1987: 136-138, pl. 39, fig. 152a, b, pl. B, fig. 4. 


Remarks 

This synonymy has long been known. The 
species, when first named, was incorrectly 
thought to have come from Africa (Hinds, 1844: 
164). Based on a petition by Bratcher & Burch 
(1971), the International Commission on Zoo- 
logical Nomenclature (1980) gave precedence 
to the better known, more appropriately named 
T. variegata for this common Panamic species 
(Keen, 1971; Bratcher & Cernohorsky, 1987). 
Holotype of 7. africana: ВММН 1872.10.12.12; 
holotype of 7. variegata: BMNH 1979121. 


Turridae 


Pleurotoma carinata 
Gray in Griffith & Pidgeon, 1833 


244 PETIT & COAN 


Pleurotoma carinata Gray in Griffith & Pidgeon, 
1833: pl. 23, fig. 2 [legend]. 
Non Pleurotome [sic] carinata Link, 1808: 36; 
non Pleurotoma decussata carinata Grate- 
leu, 18327 332. 

Pleurostoma [sic] carinata Gray — Griffith & 
Pidgeon, 1834: 599. 

Pleurotoma speciosa Reeve, 1842b: 187, pl. 
235, NG. O: 

Gemmula speciosa (Reeve, 1842) — Powell, 
1964: 47; 1966: 246. 


Remarks | 

Probably a synonym of Gemmula speciosa 
(Reeve, 1842) (Powell, 1964; 1966) from the 
Philippine Islands. BMNH 1875.4.26.1, holo- 
type of Pleurotoma carinata Gray. 


Pleurotoma grandis 
Gray in Griffith & Pidgeon, 1833 


Pleurotoma crispa Lamarck, 1816: 8, pl. 439. 

P. grandis Gray in Griffith & Pidgeon, 1833: 
pl. 23, fig. 3 [legend]. 

Pleurostoma [sic] grandis Gray - Griffith & 
Pidgeon, 1834: 599. 

Turris crispa (Lamarck, 1816) — Powell, 1964: 
33: 


Remarks 

Synonym of Turris crispa (Lamarck, 1816) 
(Powell, 1964), which occurs from Madagas- 
car to Australia and Fiji. BMNH 1875.4.29.1, 
holotype of P. grandis. 


Clavatula griffithii 
Gray in Griffith & Pidgeon, 1833 


Clavatula griffithii Gray in Griffith & Pidgeon, 
1833: pl. 37, fig. 6 [legend; first issue of pl. 
with reversed images]; 1834: pl. 37*, fig. 6 
[legend; second issue of pl. with corrected 
images]. 

С. griffithii Gray — Griffith & Pidgeon, 1834: 596. 

С. griffithii Gray — Kilburn, 1989: 189-190, figs. 
9, 10 [holotype]. 

Ptychobela griffithii (Gray in Griffith & Pidgeon, 
1833) — Hylleberg & Kilburn, 2002: 44. 

Ptychobela griffithii (Gray in Griffith & Pidgeon, 
1833) — Tucker, 2004: 441. 


Remarks 

Ptychobela griffithii (Gray in Griffith & 
Pidgeon, 1833) from the tropical western Pa- 
cific (Hylleberg & Kilburn, 2002; Tucker, 2004). 
BMNH 1875.4.26.22, holotype. 


Ferussaciidae 


Bulimus aurisvulpina 
(Holten, 1802) 


Voluta auris-vulpina Holten, 1802: 45, ex 
Chemnitz vol. Il, figs. 2086, 2087. 

Bulimus auris-vulpina Gray [sic] — Griffith & 
Pidgeon, 1834: 596 (St. Helena). 


Remarks 

Attributed to Gray in the list of figures in Griffith 
& Pidgeon (1834), an error that was picked up 
by Sherborn (1923: 597). This land snail from 
St. Helena, Chilonopsis aurisvulpinus (Holten, 
1802), is now extinct (Abbott, 1989: 84). 


Caryodidae 


Helix hayii 
Gray in Griffith & Pidgeon, 1833 


Helix hayii Gray in Griffith & Pidgeon, 1833 
[Dec.]: pl. 36, fig. 4 [legend; first issue of pl. 
with reversed images]. 

H. cunninghami Gray in Griffith & Pidgeon, 
1834: pl. 36*, fig. 1 legend; second issue of 
pl. with corrected images; March or June]. 

H. cunninghami Gray — Griffith & Pidgeon, 
1834 [June]: 597. 

H. cunninghami Gray, 1834: 64 [25 Nov.]. 

Pedinogyra hayii Griffith & Pidgeon — Iredale, 
19375415216) рт, figs. Y, 2: 


Remarks 

For reasons unknown, Gray changed his mind 
about the name of this species between the 
original issue of pl. 36 in 1833, with its reversed 
images, and the later issue of the plate with 
correct images and the list of figures in 1834, 
the only instance of such a name change. This 
species is from Queensland, Australia (Iredale, 
1937). As H. cunninghami, this is type species 
by original designation of Pedinogyra Martens, 
1860: 162-163 (Kabat & Boss, 1997: 119-120). 
BMNH 1963861, holotype. 


Camaenidae 


Helix fraseri 
Gray in Griffith & Pidgeon, 1833 


Helix fraseri Gray in Griffith & Pidgeon, 1833: 
pl. 36, fig. 6 [legend; first issue of pl. with re- 
versed images]; 1834: pl. 36*, fig. 6 [едепа; 
second issue of pl. with corrected images]. 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 245 


H. fraseri Gray — Griffith & Pidgeon, 1834: 596 
(New Holland). 


Remarks 

Type species, by the subsequent designa- 
tion of Pilsbry (1890: 84), of Sphaerospira 
Mörch, 1867: 256. Sphaerospira fraseri (Gray 
in Griffith and Pidgeon, 1833) occurs in east- 
ern Australia. BMNH 1847.4.12.17-18, 2 
syntypes. 


Helix argellacea 
Gray in Griffith & Pidgeon, 1833 


Helix argellacea Gray in Griffith & Pidgeon, 
1833: pl. 28, fig. 2 [legend; first issue of pl. 
with reversed images]; 1834: pl. 28", fig. 2 
[legend; second issue of pl. with corrected 
images]. 

H. argillacea [sic] Gray — Griffith & Pidgeon, 
1834: 597. 

Non Helix argilacea Férussac, in Ferussac 
& Deshayes, 1820: pl. 26, figs. 1-3 [figure 
without name]; 1821: 9 [pl. expl.]. 


Remarks 

Because this species name was made avail- 
able in 1833, the spelling argellacea has pre- 
cedence over the spelling argillacea a year 
later; for this reason, it is not a junior hom- 
onym of Férussac's 1821 species (ICZN Code 
Art. 58.7 — spellings differing only in a single 
or a double consonant are equivalent). While 
itis possible that the species depicted in Griffith 
& Pidgeon might have been intended to be 
that of Ferussac, the figures in the two works 
seem to be of significantly different taxa. As 
far as we know, the Gray species has not been 
subsequently recognized. Type material not 
found in 2006 in the BMNH. 


Helix mora 
Gray in Griffith & Pidgeon, 1833 


Helix carmelita Ferussac, 1820: pl. 32, fig. 4 
[three views; without name]; 1821: 10 [pl. expl.]. 
Non H. carmelita Lichtenstein, 1794: 99 [see 
Geiger, 2003: 82]. 

H. mora Gray in Griffith & Pidgeon, 1833: pl. 
36, fig. 1 [legend; first issue of pl. with reversed 
images]; 1834: pl. 36”, fig. 1 [legend; second 
issue of pl. with corrected images]; 597. 

H. carmelita Ferussac — Deshayes, 1850: 193 
[with Helix mora in synonymy]. 

Pleurodonte mora (Griffith & Pidgeon) — 
Rosenberg & Muratov, 2006: 130-131, 161. 


Remarks 

This species occurs in Jamaica. Type mate- 
rial not present in the BMNH (Rosenberg & 
Muratov (2006). 


ACKNOWLEDGEMENTS 


We appreciate the assistance of the many 
persons who have assisted us in preparing this 
paper. The staff of The Natural History Mu- 
seum allowed one of us (EVC) to search the 
collection for type material and provided ad- 
vice on particular taxa, including Joan 
Pickering, David Reid, and Kathie Way. Julie 
Gardham, Fiona Neale, and Archie Pollock, 
Glasgow University Library, correspondended 
on bibliographical matters. Rüdiger Bieler sup- 
plied a German translation. M. G. Harasewych 
and Paul Callomon provided copies of scarce 
literature. Hugh Torrens supplied biographical 
information about Edward Pidgeon. Alan Kabat 
reviewed an earlier draft of the paper, and two 
anonymous reviewers provided helpful com- 
ments. Others supplied advice about the mod- 
ern allocations of the taxa involved and related 
nomenclatural issues, including Willem Back- 
huys, Arthur Bogan, Robert Cowie, Kevin 
Cummings, Neal Evenhuis, Matthias Glau- 
brecht, Frank Kohler, Harry Lee, James 
McLean, Bruce Marshall, Timothy Pearce, Win- 
ston Ponder, Gary Rosenberg, Peter Skelton, 
and Geerat Vermeij. 


LITERATURE CITED 


ABBOTT, R. T., 1960, The genus Strombus in 
the Indo-Pacific. Indo-Pacific Mollusca, 1(2): 
33-146, incl. pls. 11-117. 

ABBOTT, К. Т., 1989, Compendium oflandshells. 
Melbourne, Florida, American Malacologists. 


240 pp. 
ABBOTT, R. T. & S. P. DANCE, 1982, Compen- 
dium of seashells. New York, E. P. Dutton. vii + 


411 pp. 

ADAMS, A., 1851, Monograph of Sphaenia, a 
genus of lamellibranch Mollusca. Proceedings 
of the Zoological Society of London, for 
1850[18](206): 86-89, pl. 10 [Plate with new 
species other than the bivalve subject of the 
paper]. 

ADAMS, A., 1855, Descriptions of thirty-nine new 
species of shells from the collection of Hugh 
Cuming, Esq. Proceedings of the Zoological 
Society of London, for 1854[22](268): 130-138, 
pl. 28. 

ADAMS, А. & L. REEVE, 1848-50, Mollusca. In: 
A. ADAMS, ed., The Zoology of the voyage of 


246 PETIT & COAN 


H.M.S. Samarang; under the command of Cap- 
tain Sir Edward Belcher, C.B., ER.A.S., F GS. 
during the years 1843-1846. London, Reeve, 
Benham & Reeve. x + 87 pp., 24 pls. [Pp. i-x 
(by Adams only), 1-24, i-ii (pl. expl.), pls. 1-9, 
1 November 1848; 25—44, pls. 10-17, 27 April 
1850; 45-87, pls. 18-24, 31 August 1850]. 

ADAMS, H., 1866, Description of a new genus 
and a new species of mollusks. Proceedings 
of the Zoological Society of London, for 
1866(1): 150-151. 

ADAMS, H. & A. ADAMS, 1853- 1858, The gen- 
era of Recent Mollusca; arranged according 
to their organization. London, John van Voorst. 
2 vols. [1, 1-256, pls. 1-32 (1853); 1, 257-484, 
2, 1-92, pls. 33-72 (1854); 2, 93-284, pls. 73- 
96 (1855): 2, 285—412, pls. 97.112 (1856); 2. 
413-540, pls. 113- 128 (1857); 2, 541-660, ols. 
129-138 (1858).] 

ALLMON, W. D., 1990, Review of the Bullia group 
(Gastropoda: Nassariidae) with comments on 
its evolution, biogeography, and phylogeny. 
Bulletins of American Paleontology, 99(135): 
1-179. 

BENSON, W. H., 1830, Description of Nova- 
culina, a new genus of fresh-water bivalves, 
inhabiting the Ganges and its branches. Glean- 
ings in Science [Calcutta], 2(14): 63-65. 

BEQUAERT, J., 1942, Random notes on Ameri- 
can Potamididae. The Nautilus, 56(1): 20-30. 

BERNARD, F. R., Y. Y. СА! 8 В. MORTON, 1993, 
Catalogue of the living marine bivalve molluscs 
of China. Hong Kong, Hong Kong University 
Press. vil + 146 pp. 

BEU, À. G. & L.A. DE ROOIJ-SCHUILING, 1983, 
Subgeneric classification of New Zealand and 
Australian species of Paphies Lesson (Bivalvia: 
Mesodesmatidae), and names for the two spe- 
cies of tuatua in New Zealand. New Zealand 
Journal of Zoology, 9(2): 211-230. 

BIVONA-BERNARDI, A., 1832, Caratteri d’un 
nuovo genere di conchiglie della famiglie delle 
columellarie del Signor de Lamarck. Effemerdi 
Scientifiche e Letterarie per la Sicilia, 2: 8-13. 

BLAINVILLE, H. M. D. de, 1824, Mollusques. Pp. 
1-392, т: Е. ©. LEVRAULT, ed., Dictionnaire des 
Sciences Naturelles, 32: 567 pp. Paris, Le 
Normant. 

BLAINVILLE, H. M. D. de, 1825-1827, Manuel 
de malacologie et de conchyliologie; ... Paris 
& Strasbourg, Levrault. 1825: vili + 648 pp, 2 
fold-out charts; 1827: pp. 649-664, 4 + 87 + 
17 pls: 

BLAINVILLE, H. M. D. de, 1826, Pintade. 
(Conch.). P. 93, in: Е. ©. LEVRAULT, ed., Diction- 
naire des Sciences Naturelles, 41: 558 pp. 
Paris, Le Normant. 

BOSCH, D. T. & E. BOSCH, 1982, Seashells of 
Oman. London, Longman Group. 206 pp. 

BOSCH, D. T., S. P. DANCE, R. G. MOOLEN- 
BEEK & P. G. OLIVER, 1995, Seashells of east- 
ern Arabia. Dubai, Motivate Publishing. 296 pp. 

BOSS, К. 4. & К. BIELER, 1991, Johannes Thiele 
and his contributions to zoology. Part 2. Genus- 
group names (Mollusca). Nemouria, 39: 77 pp. 

BOUVIER, E.L., 1887, Observations sur le genre 
Ceratoptilus créé dans la famille des cérithidés. 


Bulletin de la Société Philomathique de Paris, 
(7) 11: 36-38 [not seen]. 

BRATCHER, Т. & К. BURCH, 1971, Terebra 
variegata Gray, 1834 (Gastropoda): proposed 
preservation under the Plenary Powers. 
Z.N.(S.) 1927. Bulletin on Zoological Nomen- 
clature, 27(5-6): 255-256. 

BRATCHER, T. & W. CERNOHORSKY, 1976, 
Living terebras of the world. Melbourne, 
Florida, American Malacologists, Inc. 236 pp. 

BRODERIP, W. J., 1833a, [Descriptions of shells 
from the Cuming collection by Mr. Broderip and 
Mr. G. B. Sowerby]. Proceedings of the Zoo- 
logical Society of London, for 1832[2], (24): 


173-179. 

BRODERIP, W. J., 1833b, [... new species ex- 
hibited ... accompanied by characters by Mr. 
Broderip]. Proceedings of the ee. Soci- 
ety of London, for 183311], (1-3): 2-8. 

BRODERIP, W. Jr 1836, [... shells of the genus 
Mitra ... and one species of Conoelix, ... form- 
ing part of the collection of Mr. Cuming ...]. 
Proceedings of the Zoological Society of Lon- 
don, for 1835[3], (36): 192-198. 

BRODERIP, W. J. & С. В. SOWERBY, |, 1829, 
Observations on new or interesting Mollusca 
contained, for the most part, in the Museum of 
the Zoological Society. Zoological Journal, 
4(15): 359-376, pl. 9. 

BRONGNIART, A., 1810, Sur des terrains qui 
paraissent avoir ete formes sur l'eau douce. 
Annales du Muséum d'Histoire Naturelle, 
15(89-90): 357-405, pls. 22, 23. 

BROWN, D. S., 1994, Freshwater snails of Af- 
rica and their medical importance, 2" ed. Lon- 
don, Taylor & Francis, x + 609 pp. 

BROWN, P. À. H., 1982, London publishers and 
printers c. 1800-1870. el) The British Li- 
brary, vii + 233 pp. 

BRUGUIERE, J. G., 1789, Encyclopédie 
méthodique. ‘Histoire naturelle de vers, 1(1): 
xviii + 344 pp. Paris, Panckoucke. 

CERNOHORSKY, W. O., 1971, Indo-Pacific 
Pisaniinae (Mollusca: Gastropoda) and related 
buccinid genera. Records of the Auckland In- 
stitute and Museum, 8, 137-167. 

CERNOHORSKY, W. O., 1975, Supplementary 
notes on the taxonomy of buccinid species of 
the subfamily Pisaniinae (Mollusca: Gas- 
tropoda). Records of the Auckland Institute and 
Museum, 12: 175-211. 

CERNOHORSKY, W. O., 1976, The Mitridae of 
the world. Part I. The subfamily Mitrinae. /ndo- 
Pacific Mollusca, 3(17): 273-528, incl. pls. 248- 
466. 

CERNOHORSKY, W. O., 1981, The family 
Buccinidae Part 1: The genera Nassaria, 
Trajana and Neoteron. Monographs of Marine 
Mollusca, 2: 1-52. 

CERNOHORSKY, W. O., 1984, Systematics of 
the family Nassariidae (Mollusca: Gastropoda). 
Bulletin of the Auckland Institute and Museum, 
14: i-iv, 1-356. 

CERNOHORSKY, W. O., 1986, Ataxonomic study 
of two Indo-Pacific species of Cantharus 
(Prodotia) [Mollusca: Buccinidae]. Records of 
the Auckland Institute and Museum, 23: 59-65. 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 247 


CHEN, S.-F., 1943, Two new genera, two new 
species, and two new names of Chinese 
Melaniidae. The Nautilus, 57(1): 19-21. 

CHENU, J. C., 1843, Solen. In: J.C. CHENU, Illus- 
trations conchyliologiques ou description et fig- 
ures de toutes les coquilles connues vivantes 
et fossiles, classées suivant le systeme 
Lamarck modifié d’après les progres de la sci- 
ence, et comprenant les genres nouveaux et 
les expéces récemment découvertes, vol. 1: 8 
ols. Paris, Franck, 4 vols., 215 pp., 482 pls. [no. 
5: pl. 4, pre-11 May 1843; no. 6: pl. 1, pre-11 
May 1843; nos. 8-10: pls. 2, 3, 5, 6, pre-10 
Aug. 1843; no. 12, 13: pl. 7, 1843; nos. 18, 19: 
pl. 8, 7 Dec. 1843] 

COAN, Е. V., 1972, What's the difference?: au- 
thorship of a taxon. The Veliger, 15(1): 64-65. 

COAN, E. V. & B. ROTH, 1976, Status of the 
genus Hyalina Schumacher, 1817 (Mollusca: 
Gastropoda). Journal of Molluscan Studies, 
42(2): 217-222. 

СОАМ, E. V., P. H. VALENTICH-SCOTT & Е. К. 
BERNARD, 2000, Bivalve seashells of west- 
ern North America. Marine bivalve mollusks 
from Arctic Alaska to Baja California. Santa 
Barbara Museum of Natural History Mono- 
graphs, 2: vili + 764 pp. 

COSEL, R. von, 1993, The razor shells of the 
eastern Atlantic. Part |: Solenidae and Pharidae 
| (Bivalvia: Solenacea). Archiv für Mollusken- 
kunde, 122: 207-321. 

COUNTS, С. L., Ш, 1991, Corbicula (Bivalvia: 
Corbiculidae). Part 1. Catalogue of fossil and 
Recent nominal species. Part 2. Compendium 
of zoogeographic records of North America and 
Hawaii, 1924—1984. Tryonia, 21: ii + 134 pp. 

COWAN, C. F., 1969, Notes on Griffith’s Animal 
Kingdom of Cuvier (1824-1835). Journal of the 
Society for the Bibliography of Natural History, 
5(2): 137-140. 

COWAN, ©. F., 1971, On Guérin's /conographie: 
particularly the insects. Journal of the Society 
for the Bibliography of Natural History, 6(1): 18- 
29 


COWIE, R. H., 1997, Catalog and bibliography 
of the nonindigenous nonmarine snails and 
slugs of the Hawaiian Islands. Bishop Museum 
Occasional Papers, 50: 1-66. 

COWIE, RH & S. СОТНЕМСО, 2003: The 
apple snails of the Americas (Mollusca: Gas- 
tropoda: Ampulariidae: Asolene, Felipponea, 
Marisa, Pomacea, Pomella): a nomenclatural 
and type catalogue. Malacologia, 45(1): 41- 
100. 

CROSSE, H., 1862, Description d’une espece 
nouvelle appartenant au genre Pisania. Jour- 
nal de Conchyliologie, 10(3): 251-252, pl. 10. 

CUVIER, BARON 16.3, LAN-6F:D.], 1016, Le 
règne animal distribué d’après son organiza- 
tion, pour server de base а l'historie naturelle 
des animaux et d’introduction a l’anatomie 
compare. Vol. 2, Les reptiles, les poisons, les 
mollusques et les annéledes. Paris, Deterville. 
xviii + 532 pp. 

CUVIER, BARONHG: deb NICH FD] 1822, 
Recherches sur les ossemens fossiles, ou Гоп 
rétablit les caractéres de plusiers animaux don't 


les revolutions du globe ont détruits les espèces; 
.... Nouvelle edition, entièrenebt refiydyem et 
considéreablement augmentée, vol. 2(2), 
Contenant la description géologique des 
couches des environs de Paris, parmi lesquelles 
se trouvent les gypses à ossemens ... Paris & 
Amsterdam, С. Dufour & Е. D'Ocagne. 648 pp., 
59 pls., 2 maps. 

GUVIERFBARONTG 5. №6 F°2:4829 
1830, Le règne animal distribué d'apres son 
organisation, pour servir de base a l'histoire 
naturelle des animaux et d'introduction a 
l'anatomie comparée. Nouvelle édition, revue 
et augmentée. Paris, Déterville. 5 volumes [v. 
1: хххм + 584 pp., 1829; v. 2: xv + 406 pp. 
1829: RR  SO04spp.; 20-pls. 1830; v4: 
xxvii + 584 pp., by P. A. Latreille, 1829; v. 5: 
xxiv + 556 pp., by P. A. Latreille, 1829]. 

CUVIER, BARON [С. J. L. М. С.Е. D.], [1833]- 
37, The animal kingdom, arranged according 
to its organization, serving as a foundation for 
the natural history of animals, and as an intro- 
duction to comparative anatomy. The Crusta- 
cea, arachnides, & Insecta, by M. Latrielle, ... 
translated from the latest French Edition. Lon- 
don, G. Henderson. 4 vols. (in 8) [Collation in 
Appendix C]. 

DA COSTA, E. M., 1778, Historia naturalis 
testaceorum Britanniae, or the British conchol- 
ogy; containing the descriptions and other par- 
ticulars of natural history of the shells of Great 
Britain and Ireland: illustrated with figures. Lon- 
don, da Costa. xii + 254 + [x] pp., 17 pls. 

DALL, W. H., 1924, Notes on molluscan nomen- 
clature. Proceedings of the Biological Society 
of Washington, 37: 87-90. 

DANCE, S. P., 1972, On William Wood's Gen- 
eral Conchology and Index Testacelogicus. 
Basteria, 36(2-5): 157-162. 

DAVIS, J. D., 1965, Mesodesma deauratum: 
synonymy, holotype and type locality. The Nau- 
tilus, 78(3): 96-100, pl. 9. 

DESHAYES, G. P., 1832, Encyclopedie 
méthodique. Histoire naturelle des vers, 2(2): 
145-594. Paris, Agasse. 

DESHAYES, G. P., 1844, Histoire naturelle des 
animaux sans vertèbres, 2™ ed. Paris, J. В. 
Baillière. Vol. 10: 638 + [1] pp. 

DESHAYES, G. P., 1850, See Férussac & Des- 
hayes, 1819-1851. 

DESHAYES, G. P., 1855, Catalogue of the 
conchifera or bivalve shells of the collection of 
the British Museum. Part Il. Petricoladae (con- 
cluded); Corbiculidae. Pp. 217-292 [Date: 
Kabat, 1989]. 

DESHAYES, G. P., 1856, Descriptions de 
nouvelles espéces du genre Erycina. Proceed- 
ings of the Zoological Society of London, for 
1855[23], (295): 181-183. 

DILLWYN, L. W., 1817, A descriptive catalogue 
of Recent shells, arranged according to the 
Linnaean method; with particular attention to 
the synonymy. London, J. & A. Arch. 2 vols. 1: 
xii + 580 pp.; 2: 581-1092 + [29] pp. 

DONOVAN, E., 1824, 1825 [1822-1826], The 
naturalist’s repository, or miscellany of exotic 
natural history, exhibiting rare and beautiful 


248 PETIT & COAN 


specimens of foreign birds, insects, shells, 
quadrupeds, fishes, and marine productions, 
... London, Simpkin & Marshall. 5 vols., 180 
pls. with unnumbered pl. expl. 

FAUSTINO, L. A., 1928, Summary of Philippine 
marine and fresh water mollusks. Monographs 
of the Bureau of Science, Manila, Philippine 
Islands, 25: 1-384. 

FERUSSAC, А. Е. J. P. J. F. ФА. de, 1825, [Re- 
view of] The animal kingdom ... par Edw. 
Griffith et autres.... Bulletin des Sciences 
Naturelles et de Géologie (2 - Bulletin 
Universel des Sciences), 6: 236-239. 

FERUSSAC, A. Е. J. P. J. Е. ФА. de, 1835, [Re- 
view of] The animal kingdom ... trad. En anglais 
par Ed. Griffith ... Bulletin Zoologique, for 
1835(sec. 2): 73-74. 

FERUSSAC, AWE. P ds F. diAsde 86; P. 
DESHAYES, 1820, 1821, 1850 [1819-1851], 
Histoire naturelle générale et particuliére des 
mollusques terrestres et fluviatiles tant des 
espèces que l’on trouve aujourd’hui vivantes, 
que des dépouilles fossiles de celles qui 
n'existent plus: classes d’après les chacteres 
essentiels que presentment ces animaux et 
leurs coquilles. Paris, Baillière. 2 vols.: xvi + 
184 + 9677, 96° + 402 + 260 pp., 199 vari- 
ously numbered and unnumbered pls. + 47 + 
[4?] + [4] pp. of pls. expl. [Complete collation 
too complex for this context; pl. 26 = 26 Feb. 
1820; pl. 32 = 17 June 1820; pp. 9-10 of pl. 
expl. = 6 April 1821, all by Ferussac; Deshayes 
took over authorship in 1839; p. 193 of his text, 
18501. 

FISCHER, P., 1880-1887, Manuel de conchy- 
liologie et de paleontology conchyliologique our 
histoire naturelle des mollusques vivants et 
fossiles, ... Paris, F. Savy. xxiv + 1369 pp., 23 
pls. 

FORCELLI, D. O., 2000, Molluscos Magalla- 
nicos. Guia de moluscos de Patagonia y sur 
de Chile. Buenos Aires, Argentina, Vazquez 
Mazzini. 200 pp. 

GEIGER, D. L., 2003, A. A. H. Lichtenstein's 
(1794) Catalogus rerum naturalium rarissi- 
marum section seconds on mollusks: an ap- 
praisal of taxa described in an overlooked tome. 
Archives of Natural History, 30(1): 75-84. 

GERLACH, J., 2006, Terrestrial and freshwater 
Mollusca of the Seychelles Islands. Leiden, 
Backhuys. 141 pp., incl. 24 pls. a 

GLAUBRECHT, M., Z. FEHER & F. KOHLER, 
2007, Inventorizing an invader: annotated type 
catalogue of Corbiculidae Gray, 1847 (Bivalvia, 
Heterodonta, Veneroidea), including Old World 
limnic Corbicula, in the Natural History Museum 
Berlin. Malacologia, 49(2): 243-272. 

GMELIN, J. F., 1791, Caroli a Linne ... Systema 
naturae per regna tria naturae ... editio decima 
tertia, acuta, reformata 1(6): 3021-3910. 
Leipzig, Beer. 

GRATELOUP, J. P. S. de, 1832, Tableau (suite 
du) des coquilles fossiles qu’on rencontre dans 
les terrains calcaires tertiaires (faluns) des 
environs de Dax, département des Landes. 
Actes de la Société Linnéenne de Bordeaux, 
5(30): 314-344. 


GRAY, J. E., 1825, Alist and description of some 
species of shells not taken notice of by 
Lamarck. Annals of Philosophy, (n.s.)9[25], (2): 
134-140. 

GRAY, J. E., 1828-1830, Spicilegia zoologica; 
or original figures and short systematic descrip- 


- tions of new and unfigured animals. Issued in 


2 parts. London, Treüttel, Wurtz & Co. 1: 1-8, 
pls. 1, 2, 5 (1828); 2: 7-12, pls. 7-11 (1830) 
[plates 3, 4, and 6 were never published, but 
are present in the BMNH Library]. 

GRAY, J. E., 1831, Zoological miscellany, 1: 40 
pp., 4 pls. London, Treuttel, Wurz & Co. 

GRAY, J. E., 1834, ... various undescribed shells, 
chiefly contained in his own collection. Pro- 
ceedings of the Zoological Society of London, 
for 1834[2], (19): 63-68. 

GRAY, J. E., 1839, Molluscous animals and their 
shells. Pp. [101]-142, pls. 33-44, in: Е. W. 
BEECHEY, The zoology of Captain Beechey’s 
voyage; ... to the Pacific and Behring’s Straits 
… in 1825-1828. London, U.K., Henry С. Bohn. 
xii + 186 pp., 44 pls. [pp. 143-155 by G. B. 
Sowerby |]. 

GRAY, J. E., 1840a, Mollusca. Pp. 105-152, in: 
Synopsis of the contents of the British Museum, 
42" ed., 1% issue. London, G. Woodfall & Son. 

GRAY, J. E., 1840b, Mollusca. Pp. 86-89, 106- 
156, in: Synopsis of the contents of the British 
Museum, 42" ed., 2" issue. London, С. 
Woodfall & Son. 

GRAY, J. E., 1842, Mollusca. Pp. 48-92, in: Syn- 
opsis of the contents of the British Museum, 
44' ed. London, С. Woodfall & Son. 

GRAY, J. E., 1847, Alist of the Recent Mollusca, 
their synonyma and types. Proceedings of the 
Zoological Society of London, for 1847[15], 
(178): 129-219. 

GRAY, J. E., 1853-1857, A revision of the gen- 
era of some of the families of Conchifera or 
bivalve shells [title varies]. Annals and Maga- 
zine of Natural History, (2)11(61): 33—44 (1 Jan. 
1853); (65): 398-402 (1 May 1853); 13(77): 
408-418 (1 May 1854); 19(113): 366-373 [as 
“Part Ill. Arcadae”] (1 May 1857). 

GRAY, J. E., 1855, List of the Mollusca in the col- 
lection of the British Museum. Part 1. Volutidae. 
London, Taylor & Francis. [ii] + 23 pp. 

GRIFFITH, E. & E. PIDGEON, [1833]-1834, The 
Mollusca and Radiata. Vol. 12, in: E. GRIFFITH, 
ed., [1824]-1835, The animal kingdom ar- 
ranged in conformity with its organization, by 
the Baron Cuvier, member of the Institute of 
France, &c. &c. &c. with supplementary addi- 
tions to each order, by Edward Griffith, F.L.S., 
A.S., corresponding member of the Academy 
of Natural Sciences of Philadelphia, &c. and 
others. London, Whittaker and Co. viii + 601 
pp., 61 pls. 

GRUBER, J. W., 2004, Griffith, Edward (1790- 
1858). Oxford Dictionary of National Biogra- 
pliyy 2320355. 

GUERIN-MENEVILLE, F. E., 1829-1844, /cono- 
graphie de règne animal de С. Cuvier, ou 
représentation d'aprés nature de l’une des 
espéces les plus remarquables et souvent non 
encore figurées, de chaque genre d’animaux. 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 249 


Paris, J. B. Bailliere. 3 Vols. [Collation in Ap- 
pendix С]. | 

[GUERIN-MENEVILLE: EME ов 6: DES: 
HAYES], 1868, Les mollusques décrits et 
figurés d’après la classification de Georges 
Cuvier mise au courant des progrès de la sci- 
ence. XXXVI planches représantant en 520 fig- 
ures. Dessinées d’après nature et gravées sur 
cuivre. Les espèces les plus remarquables de 
ces animaux avec un texte descriptif. Paris, J. 
B. Baillière et fils. 74 pp., 36 plates. 

GUNTHER, A. E., 1975, A century of zoology at 
the British Museum through the lives of two 
keepers 1815-1914. London, U.K., Dawson. 
533 pp. [+ 6 pp. of Addenda & Corrigendal. 

HAAS, F., 1969, Superfamilia Unionacea. Das 
Tierreich, 88: x + 663 pp. 

HABE, T., 1977, Systematics of Mollusca in Ja- 
pan. Bivalvia and Scaphopoda. Tokyo, Hokuryu- 
kan. xiii + 372 pp. + [4] pp. errata, 72 pls. 

HANLEY, S., 1856, Index Testaceologicus, an 
illustrated catalogue of British and foreign 
Shells, ... by W. Wood ... A new and entirely 
revised edition. London, Willis & Sotheran. xx 
+ 234 pp., pls. 1-38 + suppl. pls. 1-8. 

HELBLING, С. S., 1779, Beytráge zur Kenntniss 
neuer und seltener Konchylien, aus einigen 
wienerischen Sammlungen. Abhandlungen 
einer Privatgessellschaft in Böhmen, 4: 102- 
131, 4 pls. 

HERRMANNSEN, A. N., 1846-1852, Indicis 
generum malacozoorum primordia. Casell, 
Fischer. Vol. 1: xxvii + 637 pp. [pp. i-xxvii + 1— 
232, 1846; pp. 233-637, 1847]; vol. 2: xxix—xlii 
+ 717 + + 140 pp. [pp. 1-352, 1847; 353- 
492, 1848; xxix-xlii, 493-717, 1849; Supple- 
ment, v + 140 pp., 1852]. 

HIDALGO, J. G., 1904, Catalogo de los moluscos 
testaceos de las Islas Filipinas. Jolö y Marianas. 
Livr. |. Revista de la Real Academia de Ciencias, 
Fisicas y Naturales de Madrid, 1: 1-192. [Three 
parts republished together: 1904—1905, Madrid: 
Gaceta de Madrid, xvi + 408 pp. + frontispiece 
(portrait). ] 

HIGO, S., P. CALLOMON & Y. GOTO, 1999, 
Catalogue and bibliography of the marine shell- 
bearing Mollusca of Japan. Osaka, Japan, Elle 
Scientific Publications. 749 pp. 

HINDS, R. B., 1844, Descriptions of new shells, 
collected during the voyage of the Sulphur, and 
in Mr. Cuming’s late visit to the Philippines [with] 
Synopsis of the known species of Тегерга. Pro- 
ceedings of the Zoological Society of London, 
for 1843 [11], (130): 149-168. 

HOLTEN, H. S., 1802, Enumeratio systematica 
conchyliorum beat. J.H. Chemnitzii ....Havniae, 
K. H. Scidelini. [vi] + 88 pp. 

HOUBRICK, R. S., 1981, Anatomy, biology and 
systematics of Campanile symbolicum with ref- 
erence to adaptive radiation of the Cerithiacea 
(Gastropoda: Prosobranchia). Malacologia, 
21(1-2): 263-289. 

HOUBRICK, R. S., 1984, Revision of higher taxa 
in genus Cerithidea (Mesogastropoda: 
Potamididae) based on comparative morphol- 
ogy and biological data. American Malacologi- 
cal Bulletin, 2: 1-20. 


HOUBRICK, R. S., 1986, Cerithidea reidi, spec. 
nov., from western Australia. The Veliger, 28(3): 
280-286. 

HOUBRICK, R. S., 1989, Campanile revisited: 
implications for cerithioidean phylogeny. Ameri- 
can Malacological Bulletin, 7(1): 1-6. 

HU, C.-H. & H.-J. TAO, 1995, Shells of Taiwan 
illustrated in color. Taiwan, National Museum 
of Science. v + 483 pp. 

HYLLEBERG, J. & К. М. KILBURN, 2002, Zooge- 
ography and inventory of marine molluscs en- 
countered in southern India. Phuket Marine 
Biological Center Special Publication, 26: 80 pp. 

INTERNATIONAL COMMISSION ON ZOOLOGI- 
CAL NOMENCLATURE, 1980, Opinion 1156. 
Terebra variegata Gray, 1834, (Mollusca, Gas- 
tropoda) conserved by the use of the Plenary 
Powers. Bulletin of Zoological Nomenclature, 
37(2): 94-95. 

INTERNATIONAL COMMISSION ON ZOOLOGI- 
CAL NOMENCLATURE, 1986, Opinion 1372. 
Donax hanleyanus Philippi, 1847 (Mollusca, 
Bivalvia): conserved. Bulletin of Zoological 
Nomenclature, 43(1): 21-22. 

INTERNATIONAL COMMISSION ON ZOOLOGI- 
CAL NOMENCLATURE, 1986, Opinion 1414. 
Panopea Menard de la Groye, 1807 (Mollusca, 
Bivalvia): conserved. Bulletin of Zoological 
Nomenclature, 43(3): 258-261. 

INTERNATIONAL COMMISSION ON ZOOLOGI- 
CAL NOMENCLATURE, 1999, /nternational 
Code of Zoological Nomenclature, 4" ed. Lon- 
don, 1.T.Z.N. xxix + 306 pp. 

IREDALE, T., 1917, More molluscan name- 
changes, generic and specific. Proceedings of 
the Malacological Society of London, 12(6): 
322-330. 

IREDALE, Т., 1937, A basic list of the land Mol- 
lusca of Australia. — Part Il. The Australian Zo- 
ologist, 9(1): 1-39, pls. 1-3. 

КАВАТ, А. R., 1989, The “Gray catalogues” [Mol- 
lusca] of the British Museum. The Nautilus, 
103(3): 113-115. 

KABAT, A. R. & K. J. BOSS, 1997, Karl Eduard 
von Martens (1831-1904): His life and works. 
Department of Mollusks, Museum of Compara- 
tive Zoology, Cambridge, Massachusetts. vii + 
417 pp. 

KAICHER, S. D., 1985, Nassariidae Part Ill. Card 
catalogue of world-wide shells, Pack 41: 4094— 
4199. 

KEEN, А. M., 1958, Sea shells of tropical west 
America. Stanford, California, Stanford Univer- 
sity Press. 624 pp., 10 pls. 

KEEN, A. M., 1971, Sea shells of tropical west 
America, 2" ed. Stanford, California, Stanford 
University Press. xiv + 1064 pp., 22 pls. 

KEEN, А. M. & К. CASEY, 1969, Superfamily 
Corbiculacea. Pp. 664-670, in: L.R. COX ETAL., 
Part N [Bivalvia], Mollusca 6, vols. 1 and 2: 
xxxvii + 952 pp., in: R. C. MOORE, ed., Treatise 
on invertebrate paleontology. Lawrence, Kan- 
sas, Geological Society of America & Univer- 
sity of Kansas. 

KING, P. P. & W. J. BRODERIP, 1832, Descrip- 
tion of Cirrhipedia, Conchifera and Mollusca, 
in a collection formed by the officers of H.M.S. 


250 PETIT & COAN 


Adventure and Beagle employed in surveying 
the southern coasts of South America, includ- 
ing the Straits of Magalhaens and the coast of 
Tierra del Fuego. The Zoological Journal, 
5(3)[49]: 332-349. 

KILBURN, R. N., 1989, Notes on Ptychobela and 
Brachytoma, with the description of a new spe- 
cies from Mozambique (Mollusca: Gastropoda: 
Turridae). Annals of the Natal Museum, 30: 
185-196. 

KOHLER, F. & M. GLAUBRECHT, 2001, Toward 
a systematic revision of the southeast Asian 
freshwater gastropod Brotia H. Adams, 1866 
(Cerithoidea: Pachychilidae): an account of 
species from around the South China Sea. 
Journal of Molluscan Studies, 67(3): 281-318. 

KOHLER, F. & M. GLAUBRECHT, 2002, Anno- 
tated catalogue of the nominal taxa of South- 
east Asian freshwater gastropods, family 
Pachychilidae Troschel, 1857 (Mollusca, 
Caenogastropoda, Cerithioidea), with an evalu- 
ation of the types. Mitteilungen aus dem Mu- 
seum für Naturkunde in Berlin, Zoologische 
Reihe, 78(1): 121-156. 

KOHLER, F. & M. GLAUBRECHT, 2006, A sys- 
tematic revision of the Southeast Asian fresh- 
water gastropod Brotia (Cerithioidea: 
Pachychilidae). Malacologia, 48(1-2): 159-251. 

LAMARCK, [J. B. Р.А. de M.] de, 1799, Prodrome 
d'une nouvelle classification des coquilles. 
Memoires de la Société d'Histoire Naturelle de 
Paris, 1: 63-80. 

LAMARCK, J. В. [P. A. de M.] de, 1801, Systeme 
des animaux sans vertébres. Paris, Deterville. 
vill + 432 pp. 

LAMARCK, J. B. P. A. [de M.] de, 1804, Suite 
des mémoires sur les fossiles des environs de 
Paris. Annales du Muséum National d'Histoire 
Naturelle, 4(19): 46-55 [Reprinted 1978, Pa- 
leontological Research Institution, Ithaca, New 
York]. 

LAMARCK, [J. B. P. A. de M.] de, 1816, Liste 
des objets représentés dans les planches de 
cette livraison. Part 23, 16 pp. In: Tableau 
encyclopédie et méthodique des trois regnes 
de la nature. Paris. 

LAMARCK, [J. В. Р.А. de M.] de, 1818, Histoire 
naturelle des animaux sans vertèbres, 5: 1- 
612. Paris, Verdière, Deterville, & chez l’auteur. 

LAMARCK, [J. B. Р.А. de M.] de, 1822, Histoire 
naturelle des animaux sans vertèbres, 6(2): 1- 
232. Paris, Verdière, Deterville, & chez l’auteur. 

LAMY, E., 1914, Révision des Mesodesmatidae 
vivants du Muséum d’ Histoire Naturelle de 
Paris. Journal de Conchyliologie, 62(1): 1-74, 
ple dee, 

LAMY, E., 1917, Révision des Crassatellidae vi- 
vants du Muséum d’Histoire Naturelle de Paris. 
Journal de Conchyliologie, 62(4): 197-270, pl. 
6 


LEA, 1., 1834, Observations on the naïads; and 
decriptions of new species of that and other 
families. Transactions of the American Philo- 
sophical Society, (n.s.) 5: 23-119, pls. 1-19 [Is- 
sued in five parts: Main Part: pp. 23-58; 
Supplement: 59-94; Second Supplement: 95- 


113; Third Supplement: 114-117; Systematic 
Index: 118-119; issued in Aug.-Sept. 1834; 
reprinted as volume, 1837]. 

LEA, I., 1841, New fresh water and land shells. 
Proceedings of the American Philosophical 
Society, 2(2): 11-15. 

LEA, I., 1843, Change of several specific names 
of melaniae, because of preoccupation. Pro- 
ceedings of the American Philosophical Soci- 
ety, 2(24): 237. 

LEA, |., 1868, Descriptions of four new species 
of exotic Unionidae. Proceedings of the Acad- 
emy of Natural Sciences of Philadelphia, 20: 
150. 

LEACH, W. E., 1814, The zoological miscellany; 
being descriptions of new, or interesting ani- 
mals, 1: 144 pp., 60 pls. London, Nodder & 
Son. 

LESUEUR, С. A., 1817, Characters of a new 
genus, an descriptions of three new species 
upon which it is formed; discovered in the At- 
lantic Ocean, in the months of March and April, 
1816; Lat. 22°9’. Journal of the Academy of 
Natural Sciences of Philadelphia, 1(3): 37-41, 
pl. 2. 

LI, X., 2004, Neogastropoda. Pp. 81-130, pls. 
50-77, in: 7. Ql, ed., Seashells of China. Beijing, 
China, China Ocean Press. viii + 418 pp., 193 


pls. 

LICHENSTEIN, А.А. H., 1794, Catalogus rerum 
naturalium rarissimarum, ... , section secunda 
continens conchylia, item mineralia, ligna ex- 
otica, & arte parata. Hamburg, Schniebes. 118 
pp. [Concerning: Geiger, 2003]. 

[LIGHTFOOT, J.], 1786, A catalogue of the Port- 
land Museum, lately the property of the Duch- 
ess Dowager of Portland, deceased, which will 
be sold at auction, by Mr. Skinner and Co. ... 
London. viii + 194 pp. 

LINK, H. F., 1808, Beschreibung der Naturalien- 
Sammlung der Universitat zu Rostock, Nach- 
trage und Verbesserungen. Rostock, Adlers 
Erben. 37 pp. 

LINNAEUS, C., 1758, Systema naturae per 
regna tria naturae ... editio decima, reformata 
1 [Regnum animale]. Stockholm, Sweden, L. 
Salvii. 824 + iii pp. 

LOBBECKE, C. H. W. T. & W. KOBELT, 1881- 
1886, Die Gattung Crassatella Lam. In: H. C. 
KUSTER & S. CLESSIN, eds., Systematisches 
Conchylien-Cabinet von Martini und Chemnitz, 
10(1a): 39 + [2] pp., 9 pls. [pp. 1-16, pls. 1-6, 
Löbbecke & Kobelt, 1881; pp. 17-41, pls. 7-9, 
Kobelt, 1886]. 

LUDBROOK, N. H., 1953, Systematic revision 
of the volutid genus Amoria. Proceedings of 
the Malacological Society of London, 30(4/5): 
131-153, pls. 14-18. 

MA, X., 2004, Mesogastropoda. Pp. 31-81, pls. 
14-50, in: 7. Qi, ed., Seashells of China. Beijing, 
China, China Ocean Press. viii + 418 pp., 193 
pls. 

MAKIYAMA, J., 1936, The Meisen Miocene of 
North Korea. Kyoto Imperial University, Mem- 
oirs of the College of Science, (B) 11(4): 193- 
228, pls. 4, 5. 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 251 


MANDAHL-BARTH, G., 1967, Revision of the Af- 
rican genera Potadoma Gray [sic] and Potado- 
moides Leloup and description of a new species 
of Cleopatra (Gastropoda Prosobranchia: 
Melaniidae). Revue de Zoologie et de Botanique 
Africaines, 76(1-2): 101-131, pls. 1, 2. 

MARSHALL, B. A., 1995, A revision of the Re- 
cent Calliostoma species of New Zealand (Mol- 
lusca: Gastropoda: Trochoidea). The Nautilus, 
108(4): 83-127. 

MARTENS, K. E. von, 1860, Die Heliceen, nach 
naturlicher Verwandtschaft systematisch 
geordnet von J. Chr. Albers. Zweite Ausgabe 
nach dem hinterlassenen Manuscript besorgt 
von Eduard von Martens. Leipzig, Wilhelm 
Engelmann. xvili + 359 pp. 

McMURTRIE, H., 1832, Remarks. Pp. i-vi, in: G. 
CUVIER, The animal kingdom, arranged in con- 
formity with its organization. By the Baron 
Cuvier, ... Translated from the French, and 
abridged for the use of schools, &c.... New York, 
С. & С. & Н. Carvill. м + 532 pp. 

MEGERLE VON MUHLFELD, J. K., 1811, Entwurf 
eines neuen System’s der Schaltiergehäuse. 
Magazin fur die Neuesten Entdeckungen in der 
Gesammten Naturkunde, Gesellschaft 
Naturforschender Freunde zu Berlin, 5: 38-72. 

MENARD ОЕ LArGROYE}; FA. -B.; 1807, 
Mémoire sur un nouveau genre de coquille bi- 
valve-équivalve de la famille des solenoides, 
intermédiaire aux solens et aux myes, voisin 
par conséquent des glycimères; ... Memoires 
Annales du Museum [National] d’Histoire 
Naturelle, 9(50-51): 131-139, pl. 12. 

MILLARD, V., 1997, Classification of Mollusca. 
Rhine Road, South Africa: Printed by the au- 
thor, 544 pp. 

MILLARD, V., 2004, Classification of Mollusca, 
2™ ed. Rhine Road, South Africa, Printed by 
the author. 3 vols. (1: 1-890; 2: 891-1,280; 3: 
1,281-1,817.) [Imprinted 2003, but published 
June, 2004]. 

MONTFORT, D. de, 1810, Conchyliologie 
systématique, et classification méthodique des 
coquilles. Vol. 2: Coquilles univalves, non 
cloisonnées. Paris, F. Schoel. 676 pp. 

MORCH, О. А. L., 1867, Abrégé de l'histoire de 
la classification moderne des Mollusques 
basée principalement sur l'armature linguale. 
Journal de Conchyliologie, 15(3): 232-258. 

MORRISON, J. P. E., 1954, The relationships of 
Old and New World melanians. Proceedings 
of the United States National Museum, 
103(3325): 357-394, pl. 11. 

MORRISON, J. P. E., 1971, Western Atlantic 
Donax. Proceedings of the Biological Society 
of Washington, 83(48): 545-568. 

NARCHI, W., 1983, Donax hanleyanus Philippi, 
1847, proposed conservation and suppression 
of Donax hilairea Guerin, 1832 (Mollusca, 
Bivalvia). Z.N.(S.)2152. Bulletin of Zoological 
Nomenclature, 40(3): 188. 

NARCHI, W., 1986, On the conservation of the 
binomen Donax hanleyanus Philippi, 1847. 
Universidade de Sao Paulo, Boletim de 
Zoologia, 10: 305-310. 


NARDO, G. D., 1840, Spora un nuovo genere di 
conchiglie de mare Adriatico. Annali delle 
Scienze del Regno Lombardo-Veneto, 10: 49- 
51: 

NEAVE, S. A., 1939-1940, Nomenclator Zoo- 
logicus. London, Zoological Society of London. 
4 volumes: xiv + 957 + 1025 + 1065 + 758 pp. 

NISSEN, C., 1969, Die Zoologische Buch-Illus- 
tration. Band I: Bibliographie. Stuttgart, Ger- 
many, Anton Hiersemann. 666 pp. 

OKUTANI, T., 2000, Subclass Anomalo- 
desmacea. Pp. 1032-1047, in: T. OKUTANI, ed., 
Marine mollusks in Japan. Tokyo, Japan, Tokai 
University Press. xlvili+ 1175 pp., incl. 542 pls. 

OLIVER, W. R. B., 1926, New Zealand species 
of Calliostoma. Proceedings of the Malacologi- 
cal Society of London, 17(2/3): 107-115, pl. 
10: 

PERRY, G., 1810-1811, Arcana, or the Museum 
of Natural History; containing the most recent 
discovered objects. London, James Stratford. 
84 pls. with unnumbered text. [Issued in parts, 
pls. 1-48 in 1810; pls. 49-84 in 1811]. 

PETIT, R. E., 1987, A final note on Cancellaria 
nassiformis Lesson, 1842, and Nassarius 
corpulentus (C. B. Adams, 1852). The Veliger, 
29(3): 340. 

PETIT, R. E., 2006, Notes on Sowerby’s The 
genera of recent and fossil shells (1821-1834). 
Archives of Natural History, 33(1): 71-89. 

PETIT, R. E., 2007, Lovell Augustus Reeve 
(1814-1865): his businesses and publications. 
Zootaxa, 1648: 120 pp. 

PETIT DE LA SAUSSAYE, S., 1853, Notice sur 
le genre Phos de Denys de Montfort, avec la 
description de deux espèces nouvelles. Jour- 
nal de Conchyliologie, 4(3): 235-245, pl. 8. 

PHILIPPI, R. A., 1850 [1846-1855], Die Kreisel- 
schnecken oder Trochoideen (Gattungen 
Turbo, Trochus, Solarium, Rotella, Delphinula, 
Phasianella). Abbildungen nach der Natur mit 
Beschreibungen. In: H.C. KUSTER & $. CLESSIN, 
eds., Systematisches Conchlien-Cabinet von 
Martini und Chemnitz, 2(3): 372 pp., 49 pls. 
[pp. 1-24, pls. 1-6, 1846; 25-32, 34, 35, pls. 
7-18, 1847; 33-72, pls. 19-24, 1848; 73-120, 
pls. 25-32, 36-39, 1849; 121-136, pls. 40-45, 
1850; 137-232, pls. 33, 46-49, 1851; 233-248, 
1852; 249-372, 1855]. 

PHILIPPI, В. A., 1847a, Versuch einer systema- 
tischen Eintheilung des Geschlechtes Trochus. 
Zeitschrift fur Malakozoologie, 4(2): 17-24. 

PHILIPPI, R. A., 1847b, Testaceorum novorum 
centuria [second of five sections]. Zeitschrift 
für Malakozoologie, 4(5): 84-96. 

PHILIPPI, R. A., 1848, Ueber die neuen Gat- 
tungen und Arten der Conchylien welche in 
Griffith’s englischer Ausgabe von Cuvier's 
Thierreich vorkommen. Zeitung für Zoologie, 
Zootomie und Palaeozoologie, 1(11): 85-87. 

PISOT DE-LAPEYROUSE, РВОТА, 
Déscription de plusieurs nouvelles espèces 
d’orthocératites de d’ostracites. De novis 
quibusdam orthoceratitum et ostracitum 
speciebus dissertatiuncula. Paris & Toulouse, 
A. Erlang. [2] + 45 + [3] pp., 13 sheets of pls. 


252 PETIT & COAN 


PILSBRY, H. A., 1889, Trochidae, Stomatiidae, 
Pleurotomariidae, Haliotidae. Manual of Con- 
chology, 11: 1-519, pls. 1-67. 

PILSBRY, H. A., 1890, [Helicidae]. Manual of 
Conchology, (2) 6(22): 65-128, pls. 16-30. 
PILSBRY, H. A. & A. HARBISON, 1933, Notes 
on the Miocene of southern New Jersey. Pro- 
ceedings of the Academy of Natural Sciences 

of Philadelphia, 85: 107-120, pls. 3-5. 

PILSBRY, Н.А. & А. А. OLSSON, 1954, Systems 
of the Volutidae. Bulletins of American Paleon- 
tology, 35(152): 270-306, pls. 25-28 [Dual pagi- 
nation; also numbered: 1-36, pls. 1-4.] 

POLI, G. S., 1791, Testacea Utriusque Siciliae 
eorumque historia et anatome tabulis aeneis 
illustrata ... Parmae, ex Regio. 1: [iv] + x + 90 + 
50 + 1 + 1-74 + 1 + Ixxiii pp., 18 pls. 

POUTIERS, J.-M. & F. R. BERNARD, 1995, 
Carnivorous bivalve molluscs (Anomalo- 
desmata) from the tropical western Pacific 
Ocean, with a proposed classification and a 
catalogue of Recent species. Mémoires du 
Museum National d'Histoire Naturelle (Paris), 
167: 107-187. 

POWELL, A. W. В., 1964, The family Turridae in 
the Indo-Pacific. Part 1. The subfamily Turrinae. 
Indo-Pacific Mollusca, 1(5): 227-346, pls. 1- 
23: 

POWELL, À. W. B., 1966, The molluscan fami- 
lies Speightiidae and Turridae. Bulletin of the 
Auckland Institute and Museum, 5: 1-184, pls. 
1-23. 

POWELL, A. W. B., 1979, New Zealand Mollusca. 
Auckland, Collins. xiv + 500 pp. + 82 pls. 

PRASHAD, B., 1932, The Lamellibranchia of the 
Siboga Expedition. Systematic part II: 
Pelecypoda (exclusive of the Pectinidae). In: 
M. WEBER, ed., Siboga-Expeditie, 34(53c)[= livr. 
118]: 353 pp., 9 pls., 1 chart. Leiden, Brill. 

QI, Z., ed., 2004, Seashells of China. Beijing, 
China, China Ocean Press. viii + 418 pp., 193 
pls. 

QUOY, J. R. С. & J. P. GAIMARD, 1824-1826, 
Voyage autour du monde ... execute sur 
PUranie et la Physicienne pendant ... 1817-20 
... part M. L. de Freycinet … Zoologie. Paris. iv 
+.712 Pp... 96 pls: 

RAFINESQUE, С. S., 1833, On 5 new fresh wa- 
ter shells, of Bengal and Assam in Asia. Atlan- 
tic Journal and Friend of Knowledge; a 
Quarterly Journal of Historical and Natural Sci- 
ences, Useful Knowledge, &c. with Figures, 5: 
165-166 [Spring 1833]. 

REEVE, L. [A.], 1842a, Monograph of 
Crassatella, a genus of acephalous mollusks 
(family Mactracea). Proceedings of the Zoo- 
logical Society of London, for 1842[10], (109): 
42-46 [Reprinted in Annals and Magazine of 
Natural History, (1)11(70): 302-306, 1843]. 

REEVE, L. [A.], 1842b, Conchologia systematica, 
or complete system of conchology; .... London, 
Longman, Brown, Green & Longman’s. Vol. 2: 
pp. 1-337, pls. 130-337. 

REEVE, L. [A.], 1843, Monograph of the genus 
Crassatella. Conchologia iconica; or, illustra- 


tions of the shells of molluscous animals, 1: 3 
pls. with unpaginated text. 

REEVE, L. [A.], 1843-1846, Monograph of the 
genus Pleurotoma. Conchologia iconica; or, il- 
lustrations of the shells of molluscous animals, 
1: 40 pls. [pls. 1-18, 1843; 19, 1844; 20-33, 
1845; 33-40, 1846]. 

REEVE, L. [A.], 1844, Monograph of the genus 
Triton. Conchologia iconica; or, illustrations of 
the shells of molluscous animals, 2: 20 pls. with 
unpaginated text. 

REEVE, L. [A.], 1845, Monograph of the genus 
Isocardia. Conchologia iconica; or, illustrations 
of the shells of molluscous animals, 2: 1 pl. 

REEVE, |. [A.], 1846-1847, Monograph of the 
genus Buccinum. Conchologia iconica; or, il- 
lustrations of the shells of molluscous animals, 
3: 14 pls. with unpaginated text [pls. 1-12, 
1846; 13, 14, 1847]. 

REEVE, L. [A.], 1847a, Monograph of the genus 
Pyrula. Conchologia iconica; or, illustrations of 
the shells of molluscous animals, 4: 9 pls. with 
unpaginated text. 

REEVE, L. [A.], 1847b, Monograph of the genus 
Paludomus. Conchologia iconica; or, illustra- 
tions of the shells of molluscous animals, 4: 1 


pl. 

REEVE, L. [A.], 1849, Monograph of the genus 
Eglesia. Conchologia iconica; or, illustrations 
of the shells of molluscous animals, 5: 1 pl. 

REEVE, L. [A.], 1859-1861, Monograph of the 
genus Melania. Conchologia iconica; or, illus- 
trations of the shells of molluscous animals, 
12: 59 pls. [pls. 2-17, 1859; 18-49, 1860; 1, 
50-59, 1861]. 

REID, D. G, 1986, The littorinid molluscs of 
mangrove forests in the Indo-Pacific region. 
London, British Museum (Natural History). xv 
+ 228 pp. 

REID, D. G., 1999, The genus Littoraria Griffith 
& Pidgeon, 1834 (Gastropoda: Littorinidae) in 
the tropical eastern Pacific. The Veliger, 42(1): 
21-53. 

RISSO, A., 1826, Histoire naturelle des 
principales productions de l’Europe méri- 
dionale et principalement de celles des envi- 
rons de Nice et des Alpes-Maritimes. 
Mollusques. Paris, Levrault. Vol. 4: vii + 439 
Spat2 pls: 

ROBBA, E., 1. DI GERONIMO, М. СНАМЕМЕЕ, 
M. P. NEGRI & КВ. SANFILIPPO, 2002, Ho- 
locene and Recent shallow soft-bottom mol- 
lusks from the northern Gulf of Thailand area: 
Bivalvia. Bollettino Malacologico, 38(5-8): 49- 
Не pls: 

ROBINEAU-DESVOIDY, A. J. B., 1830, Essai sur 
les myodaires. Mémoires Présentés par Divers 
Savans a l’Académie Royale des Sciences de 
l'Institut de France, ..., Sciences Mathé- 
matiques et Physiques, 2: 813 pp. 

RODING, P. F., 1798, Museum Boltenianum sive 
catalogus cimeliorum e tribus regnis naturae 
quae olim collegerat Joa. Fried Bolten, M.D.p.d. 
per XL. annos proto physicus hamburgensis. 
Pars secunda contiens conchylia sive testacea 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 255 


univalvia, bivalvia & multivalvia. Hamburg, J. 
C. Trappii. vili + 199 pp. 

ROSENBERG, С. & I. V. MURATOV, 2006, Sta- 
tus report on the terrestrial Mollusca of Ja- 
maica. Proceedings ofthe Academy of Natural 
Sciences of Philadelphia, 155: 117-161. 

SABELLI, B., R. GIANNUZZI-SAVELLI & D. 
BEDULLI, 1990, Catalogo annotato dei 
molluschi marini del Mediterraneo. Annotated 
check-list of Mediterranean marine mollusks. 
Vol. 1: xiv + 348 pp. Bologna, Italy, Libreria 
Naturalistica Bolognese. 

SAY, T., 1821, Descriptions of univalve shells of 
the United States. Journal of the Academy of 
Natural Sciences of Philadelphia, 2(1): 149-178. 

SAY, T., 1822, An account of some of the marine 
shells of the United States — concluded. Jour- 
nal of the Academy of Natural Sciences of 
Philadelphia, 2: 302-325. 

SCHUMACHER, C. F., 1817, Essai d’un nouveau 
Système des habitations des vers testacés. 
Copenhagen, Denmark, Schultz. 287 pp., 22 
pls. 

SEGUENZA, L., 1903, Molluschi poco noti dei 
terreni terziarii di Messina. Trochidae e 
Solariidae. Bollettino della Societa Geologica 
Italiana, 21: 455-464. [Imprinted 1902, but 
published 1903 fide Neave, 1940, 4: 575]. 

SHERBORN, C. D., 1922-1932, Index Ani- 
malium sive index nominum quae ab A.D. 
MDCCLVIII generibus et specibus animalium 
imposita sunt. Section Secunda. A kalendis 
lanuariis, MDCCCI usque ad finem Decembris, 
MDCCCL. British Museum (Natural History), 
London. [Issued in 33 parts: 1, [il-exxxii, 1- 
128, 1922; 2, 129-384, cxxxili-cxxxvi, 1923; 
3, 385-640, 1923; 4, 641-943, 1924; 5, 945- 
1196, 1924; 6, 1197-1452, 1925; 7, [cxxxvii]- 
cxxxix, 1925; 8, 1773-2008, 1925; 9, 2009-2248, 
1926; 10, 2249-2568, 1926; 11, 2569-2880, 
1926; 12, 2881-3136, 1927: 13, 3137-3392, 
1927; 14, 3393-3746, 1927; 15, 3747-3970, 
1928; 16, 3971-4194, 1928; 17, 4195-4450, 
1928; 18, 4451-4690, 1929; 19, 4691-4930, 
1929; 20, 4931-5138, 1929; 21, 5139-5348, 
1929; 22, 5349-5701, 1930; 23, 5703-5910, 
1930; 24, 5911-6118, 1930; 25, 6119-6358, 
1931; 26, 6359-6582, 1931; 27, 6583-6806, 
1931; 28, 6807-7056, 1932; 29, [i]-vii, cxxxiii- 
cxlvili, 1-208, 1932; 30, 209-416, 1932; 31, 
417-654, 1932; 32, 655-878, 1933; 33, 879- 
1098, 1933] 

SIMPSON, C. T., 1900, Synopsis of the naiads, 
or pearly fresh-water mussels. Proceedings of 
the United States National Museum, 22: 501- 
1044, pl. 18. 

SMITH, J. C., 1993, Georges Cuvier. An anno- 
tated bibliography of his published works. 
Washington, D.C., & London, Smithsonian In- 
stitution. xx + 251 pp. 

SOWERBY, С. B., I, 1830, Genus Ancillaria. In: 
W. J. BRODERIP, ed., Species conchyliorum, or 
concise original descriptions and observations 
... Of all of the species of Recent shells, 1(1): 
10 pp., 3 unnumbered pls. London, Sowerby. 


SOWERBY, С. B., |, 1832a, Littorina. 2 pp., 1 pl. 
In: The genera of recent and fossil shells, for 
the use of students in conchology and geol- 
ogy, with original plates ... London, Sowerby. 
265 unnumbered plates and text [Collated in 
Petit, 2006]. | 

SOWERBY, С. B., I, 1832b, ... new species of 
shells collected by Mr. Cuming on the western 
coast of South America and among the islands 
of the South Pacific Ocean. Proceedings ofthe 
Zoological Society of London, for 1832[2], (19): 
113-120. 

SOWERBY,GeBr-l, 18332], Fusus: Spp., 1 pl. 
in: The genera of recent and fossil shells, for 
the use of students in conchology and geol- 
ogy, with original plates ... London, Sowerby. 
265 unnumbered plates and text [Collated in 
Petit, 2006]. 

SOWERBY, С. B., Il, 1859, Monograph of the 
genus Nassaria Link. In: G. В. SOWERBY, Il, ed., 
Thesaurus conchyliorum, 3: 85-87, pl. 220. 

SOWERBY, С. B., Ш, 1909, Notes on the family 
Ampullariidae, with list of species, varieties, 
and synonyms, also descriptions of four new 
species. Proceedings of the Malacologial So- 
ciety of London, 8(6): 345-362. 

STIMPSON, W., 1851, Shells of New England. A 
revision of the synonymy of the testaceous mol- 
lusks of New England. Boston, Massachusetts, 
Phillips, Sampson & Co. vi + 7-58 pp., 2 pls. 

SWAINSON, W., 1840, A treatise on malacology, 
or shells and shell-fish. London, Longman, 
Orme, Brown, Green, & Longmans. viii + 419 


pp. 

S1WENNEN: 6. Во@. MOOLENBECK, В. 
RUTTANADAKUL, H. HOBBELINK, H. 
DEKKER 8 S. HAJISAMAE, 2001, The mol- 
luscs of the southern Gulf of Thailand. 
Bangcock, Thailand, The Thailand Research 
Fund, Thai Studies in Biodiversity 4: ix + 211 
pp., incl. [44] pls. 

THIELE, J., 1929, Handbuch der systematischen 
Weichtierkunde, 1(1): 1-376. Jena, G. Fischer. 

TOPHAM, J. R., 2004. Whittaker, George Byrom 
(1793-1847), publisher and bookseller. Oxford 
Dictionary of National Biography, Vol. 5: 754- 
755. Oxford, Oxford University Press. 

TRYON, G. W., 1881, Tritonidae, Fusidae, 
Buccinidae. Manual of Conchology, 4: 1-276, 
pls. 1-58. 

TSUCHIYA, K., 2000, Family Muricidae. Pp. 365- 
421, in: T. OKUTANI, ed., Marine mollusks in 
Japan. Tokyo, Japan, Tokai University Press. 
xIvili + 1175 pp., incl. 542 pls. 

TUCKER, J. K., 2004, Catalog of Recent and 
fossil turrids (Mollusca: Gastropoda). Zootaxa, 
682: 1-1295. 

TURNER, H., 1993, Ungewöhnlich und neue 
Mitroidea aus dem Indopazifik (Teil 2). Club 
Conchylia Informationen, 25(2): 82-111, incl. 
pls. 1-4. 

TURTON, W., 1822, Conchylia Insularum 
Britannicarum. The shells ofthe British Islands, 
systematically arranged. Exeter, Collum. xlvii 
+279 pp., 20 pls. 


254 PETIT & COAN 


VAUGHT, K. C., 1989, A classification of the liv- 
ing Mollusca. Melbourne, Florida, American 
Malacologists. xii + 195 pp. 

VERMEIL, С. J., 2001, Taxonomy, distribution, 
and characters of pre-Oligocene members of 
the Cantharus group of Pisaninae (Neo- 
gastropoda: Buccinoidea). Journal of Paleon- 
tology, 75(2), 295-309. 

VERMEIL, С. J., 2006, The Cantharus group of 
Pisaniine buccinid gastropods: review of the 
Oligocene to Recent genera and description 
of some new species of Gemophos and 
Hesperisternia. Cainozoic Research, 4(1-2): 
71-96. 

VOKES, H. E., 1980, Genera of the Bivalvia: a 
systematic and bibliographic catalogue (re- 
vised and updated). Ithaca, New York, Pale- 
ontological Research Institution. xxvii + 307 pp. 

WEAVER, C. S. & J. E. duPONT, 1970, Living 
Volutes. Delaware Museum of Natural History, 
Monograph, 1: i-xv, 1-375. 

WATTERS, G. T., 2006, The Caribbean land snail 
family Annulariidae: a revision of higher taxa 
and a catalogue of the species. Leiden, Neth- 
erlands, Backhuys Publishing. [viii] + 557 + [9] 
app: 

WILKINS, С. L., 1957, The Cracherode collec- 
tion. Bulletin of the British Museum (Natural 
History), Historical Series, 1(4): 121-184, pls. 
20-25. 

WILSON, B. R., 1993-1994, Australian marine 
Shells. Kallaroo, Australia: Odyssey Publish- 
ing. 2 Vols., 1: 408 pp., 1993; 2: 370 pp., 1994. 

WOOD, W., 1828a, Index testaceologicus; or, a 
catalogue of shells, British and foreign, ar- 
ranged according to the Linnean system; with 
the Latin and English names, references to fig- 
ures, and places where found, 2" ed. [actually 
3]. London, [Wood]: int] +9212 pp 38 
pls. 

WOOD, W., 1828b, Supplement to the Index 
Testaceologicus; or a catalogue of shells, Brit- 
ish and foreign. London, [Wood]. iv + [1] + 59 
pp., 8 pls. 

WOODWARD, B. B., ed., 1903-1914, Catalogue 
of the books, manuscripts, maps and drawings 
in the British Museum (Natural History). Lon- 
don, United Kingdom, Longmans, British Mu- 
seum (Natural History). viii + 2,403 pp. [1: i-viii 
+ 1-500, 1903; 2: 501-1038, 1904; 3: 1039- 
1494, 1910; 4: 1495-1956, 1913; 1957-2403, 
19451. 

WU, W.-L., 2004, The Taiwan malacofauna. V. 
Bivalvia. Taipei, Taiwan Council of Agriculture, 
Taiwan Forest Bureau). viii + 217 pp. 

XU, F., 2004, Pholadomyoidea. Pp. 326-332, pls. 
177-178, in: Z. Ql, ed., Seashells of China. 
Beijing, China, China Ocean Press. viii + 418 
pp., 193 pls. 

YEN, T. C., 1942, A review of Chinese gastro- 
pods in the British Museum. Proceedings of 
the Malacological Society of London, 24(5-6): 
170-289, pls. 11-28. 


Revised ms. accepted 15 October 2007 


APPENDIX A 


Latinizations and Spelling errors in Griffith & 
Pidgeon not involving Gray Taxa 


À few names, mostly misspellings or im- 
proper latinization, in the text of Griffith & 
Pidgeon have been attributed to them as au- 
thors. Although most are invalid and have been 
cited as misspellings, others have been treated 
as available names in some nomenclators. 
Neave (1939-1940), for reasons unknown, 
attributed some names in the text of Griffith & 
Pidgeon to Griffith only, but attributed others 
to “Griffith & Pidgeon”, whereas names that 
appeared only on the plates or in the Index 
were attributed by him to Gray. 


Calceolaria Griffith & Pidgeon, 1833: 92 — п 
Cuvier (1830: 120), this was a vernacular, “Les 
Calceoles”. Latinized here, it is thus an avail- 
able genus-group name with only a descrip- 
tive phrase and without included species. 
Listed by Neave (1939: 523) as incertae cedis. 
It could perhaps be regarded as a nomen du- 
bium within the bivalve order Hippuritoida. 


Chicoracea Griffith & Pidgeon, 1833: 79 — 
Listed by Griffith & Pidgeon as Chicoracea 
Montf., this is a simple misspelling of Chico- 
reus Montfort, 1810: 610. Listed by Neave 
(1939: 690) as “ex Montfort’. Not defined by 
Neave, his use of “ex” is taken to indicate an 
intentional emendation as opposed to his use 
of “err. pro” for misspellings, but there is no 
evidence that the Griffith & Pidgeon spelling 
was anything but a typographical error. 


Demarestia Griffith & Pidgeon, 1833: 51 — A 
typographical error in the translation of a foot- 
note in Cuvier (1830: 69), rendered the spe- 
cies name Firoloida Demarestia as “Firoloida, 
Demarestia”, as if it were a genus-group name. 
The specific name had been capitalized by 
Cuvier, a standard practice of the time when 
citing eponyms. Properly ignored by Sherborn 
and by Neave, “Demarestia Griffith & Pidgeon, 
1834”, has been incorrectly listed as a synonym 
of the heteropod genus Firoloida Lesueur, 1817: 
38, in some nomenclators, such as Vaught 
(1989: 35) and Millard (1997: 126; 2004: 236). 


Lyra Griffith, 1834: 234 — Sherborn (1927: 
3743) repeated Herrmannsen’s (1852: 78) list- 
ing of this name as “= Harpa.” Listed by Neave 
(1939: 1021) without comment, this was sim- 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 255 


ply a name in a string of uncapitalized uninom- 
inals, some Latin genera and some vernacu- 
lar terms, and was thus not intended to 
represent a Latin genus. See additional note 
about this string of names under Vis below. 


Meleager Griffith & Pidgeon, 1833: 56 — In 
Cuvier (1830: 76) this appeared as “les 
Meleagres, Montf.”. The latinization by Griffith 
& Pidgeon (1833: 56) was accompanied by the 
descriptive term “umbilicated” and by a single 
species name, Turbo pica Linnaeus, 1758: 763. 
Montfort (1810: 206) introduced the Latin ge- 
neric name Meleagris. Although treated as a 
valid name by Neave (1940: 92), the Griffith & 
Pidgeon usage must be taken as a subsequent 
misspelling. Meleagris Montfort is a junior hom- 
onym of Meleagris Linnaeus, 1758: 156, a ge- 
nus of bird. The junior synonym in current use 
is Cittarium Philippi, 1847a: 21, a replacement 
name for Meleagris Montfort. 


Mytiloidea Brongniart in Griffith, 1834: 105 
— Attributed to Brongniart in Griffith [& Pidgeon] 
by Neave (1940: 255), this is merely a spell- 
ing error for Mytiloides Brongniart in Cuvier, 
1622820! 


Orbitulites Griffith & Pidgeon, 1833: 12 — Listed 
by Neave (1940: 451) as “pro” Orbulites 
Lamarck, 1801: 451. The distinction made by 
Neave between “ex” and “pro” (see under Chi- 
coracea above) is unknown, but this is a simple 
spelling error, not intended to be a new name. 


Ostracita Griffith & Pidgeon, 1833: 92 — This 
name appeared in Cuvier (1830: 119) as the 
vernacular “Ostracite” attributed to La Peyrouse 
[sic], which was Latinized as Ostracita by Griffith 
& Pidgeon (1833: 92), here a genus without 
species. Although Neave (1940: 483) listed as 
“Ostracita Griffith 1834, in Cuvier”, it is here 
considered to be a misspelling of Ostracites 
Lamarck, 1799: 81, who named it without ref- 
erence to any species or figure. From its place- 
ment in Lamarck, it is a nomen dubium within 
the bivalve order Hippuritoida. However, this 
name is possibly attributable to Picot de Lapei- 
rouse (1781), whose work we have not seen. 


Pentadina Gray in Griffith & Pidgeon, 1834: 
599. Listed by Sherborn (1929: 4838) and by 
Neave (1940: 651) as an available name with- 
out comment, this is a misspelling of Pintadina 
Blainville, 1826: 93, with the correct spelling 
on Griffith & Pidgeon’s plate 39. 


Potamis Gray in Griffith & Pidgeon, 1833: 
pl. 32; 1834: 599. Listed by Sherborn (1929: 
4838) and by Neave (1940: 879) as a valid 
name without comment, this is a misspelling 
of Potamides Brongniart, 1810: 368. 


Vis Griffith & Pidgeon, 1834: 197. This name 
appears in Griffith & Pidgeon only in the Supple- 
ment where few genera are mentioned by 
name, and then with the initial letter not capi- 
talized and often not in italics. The text contain- 
ing this putative name, a discussion of features 
of the gastropod proboscis is, with capitaliza- 
tion and italics as printed: “This takes place in 
buccinum and neighbouring genera. In the vis 
maculata, which also has a very long probos- 
cis, ...”. While this is a binomin, the “genus” is 
not capitalized, but neither is “buccinum” capi- 
talized. Arguing against acceptance as an avail- 
able name introduction is the fact that the 
species cannot be identified without knowing 
that vis is the French vernacular term for spe- 
cies of Terebra. Although Sherborn (1932: 
6952) listed Vis because Herrmannsen (1849: 
696) had done so, he qualified it with “sed usu 
gen.?” Vis was also listed by Neave (1940: 643) 
as a nomen nudum. That conclusion was pos- 
sibly the result of assuming that vis should have 
been capitalized in the “binomin,” and that the 
species cannot be determined. We here con- 
sider it a usage of the French vernacular not 
entering nomenclature even as a nude name. 
The name appeared again in Griffith & Pidgeon, 
on page 234. There “vis” is one of a string of 
names, termed “genera,” all uncapitalized, non 
italicized, some of which are Latin (pleurotoma, 
tonna), whereas others are vernacular (cones, 
porcelaines). 


There are several typographical errors on the 
plates and Index to Plates in Griffith & Pidgeon 
that do not involve Gray taxa: 


Annatina — legend on pl. 22 — Spelling error 
for Anatina Lamarck, 1818: 462, non 
Schumacher, 1817: 125; listed by Neave as 
an error for Anatina Schumacher, 1817. 


Chonarus — legend to pl. 27 and Chrondrus — 
р. 596- Spelling errors for Chondrus Cuvier, 
1816: 408. 


Cytherea dronea — legend on pl. 19 and 
Cytherea dronia — p. 597 — Spelling errors 
for Cytherea dione (Linnaeus, 1758: 684 — 
originally as Venus). 


256 PETIT & COAN 


Pleurostoma - р. 599 — Spelling error for 
Pleurotoma Lamarck, 1799: 73; rendered 
correctly on Plate 23. 


Other spelling errors appear only in the text, 
which is in no way associated with Gray. 
Those errors listed by Neave, noted by him 
as “pro” or “err. pro” (with correct spelling in 
parentheses) are: 


Anula, р. 70 (Ovula Bruguiére, 1789: xv) 

Aspergillium, p. 125 (Aspergillum Lamarck, 
1818: 429) | 

Aximea, p. 103 (Axinaea Poli, 1791: 32) 

Brontis, p. 79 (Brontes Montfort, 1810: 622) 

Dipsada, p. 106 (Dipsas Leach, 1814: 119) 

Plocameros, p. 37 (Plocamoceros Cuvier, 
1830: 52) 

Sepiotheuthes, p. 11 (Sepioteuthis Blainville, 
1824: 175) 

Stromatia, p. 86 (Stomatia Helbling, 1779: 124) 

Timorienna, р. 52 (Timoriena Quoy 8 
Gaimard, 1824: 493) 

Trophona, p. 79 (Trophon Montfort, 1810: 482) 

Velata, p. 64 (Velates Montfort, 1810: 354) 


Although involving neither spelling nor lat- 
inization, the engraver who copied the Guérin- 
Méneville plates for Griffith 4 Pidgeon made a 
singular error. The centerpiece of Guérin- 
Méneville's plate 17 (fig. 5) is a living Buccinum 
laevissimum Gmelin, 1791: 3494. Among 
smaller shells on that same plate is Nassa 
reticulata Link, 1807: 13 (fig. 6). This large fig- 
ure of Buccinum laevissimum was among fig- 
ures reproduced in reduced size in Griffith & 
Pidgeon, where it became figure 6 on plate 32. 
The engraver did a credible job of reproducing 
the figure in a smaller size, but he copied the 
name of figure 6, Nassa reticulata, a species 
that does not appear in Griffith & Pidgeon. This 
figure is also listed in Griffith & Pidgeon’s Index 
(p. 598) as Nassa reticulata, With the further 
indignity of having an incorrect plate number 
(22) assigned. This is certainly evidence that 
Gray did not compose the entire Index, because 
he would not have confused these two species. 


APPENDIX B 
Notes on Wood's Index Testaceologicus 


Wood's works (1828а, b) can be a source of 
confusion for those who have not worked with 
them extensively. Here we provide additional 
detail about format of these books. Moreover, 
Hanley (1856) published a “revised edition” in 


which he recognized the source of many fig- 
ures and attributed authorship. It is unfortu- 
nate that he neither explained Wood's unique 
arrangement nor provided an index. 

Wood (1828a) is a list of known species, with 
figures, arranged and classified by the Lin- 
naean system. It was his intent (Wood, 1828a: 
iii) to “incorporate in one volume figures of all 
the known shells, reduced indeed to a small 
size, .... The format was described by Wood 
(1828a: v) as: “Each page is divided into five 
columns. The first contains the Linnean names: 
the second, a reference to the page in Gmelin’s 
enlarged edition of the Systema Naturae, 
where the descriptions of such shells as were 
noticed by that editor will be found. The En- 
glish names occupy the third column; the fourth 
refers to some of the principal authors whose 
figures have an established reputation; and in 
the fifth division will be seen the name of the 
place where every shell is to be found.” 

Wood also gave a list (pp. xv-xx) of 
“Lamarck’s divisions of the Linnean genera, 
referred to the figures of the Index Testa- 
ceologicus.” The list does not include all fig- 
ures, and the Lamarckian genera are not 
mentioned elsewhere in 1828a. 

Wood did not mention the very first column, 
which is a figure number. Each of the 38 plates 
contains at least 60 small figures. When a plate 
contains species of more than one genus, it is 
divided into sections delineated by a solid line, 
with the genus name in each section. Thus, to 
cite the first figure, Chiton squamosus, it does 
not suffice to reference plate 1, figure 1, as 
there is another figure 1 on the same plate, a 
species of Lepas. The only practical citation 
of the page, plate, and figure for this Chiton 
species is thus: Chiton squamosus, р. 1, pl. 1, 
Chiton fig. 1. 

The real problems arise in deciphering and 
citing the “Supplement” (Wood, 1828b), which 
contains eight additional plates numbered sim- 
ply 1 to 8. In the Hanley edition of 1856, the 
eight Supplemental plates have “Sup.” added 
after the plate number, such identification not 
being necessary earlier as the Supplement 
was originally published as a separate work. 
The Supplement is composed of, in addition 
to the Preface and plates, two sections. The 
first section (pp. 1-27) is arranged exactly as 
in 1828a. The second section (pp. 29-59) is 
headed: “References from Lamarck’s Animaux 
sans Vertebres, adapted to the figures in the 
Index Testaceologicus.” That heading is fol- 
lowing by: “The names in Italics refer to the 
Supplement, and are not in Lamarck.” The La- 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 291 


marckian genera, arranged alphabetically, are 
written in all capital letters. Below each genus 
is a reference to where coverage of the genus 
begins in Lamarck (e.g., “ACESTA, Vol. 5, p. 
Sa 

The listings are arranged in five columns, 
the first being the number of the species in 
Lamarck. The species first listed are those that 
appear in Lamarck and to which Linnaean 
genera were applied in 1828a. Next are listed, 
in italics without a number, those species 
placed in Linnaean genera on pages 1-27 of 
1828b. 

The second column is the species name (not 
italicized for nomina appearing in 1828a). 
Forming a third column is the Lamarckian ge- 
nus to be applied, italicized and in parenthe- 
ses. It takes study and comparison to realize 
that the absence of a Lamarckian genus indi- 
cates that it the same as the prior listing, as 
there are no solid lines to indicate “ditto marks” 
as in 1828a. 

The final two columns are the plate and fig- 
ure numbers that apply. Of course, plate and 
figure references for non-italicized species are 
to 1828a. In the Hanley edition, the listings for 
the Supplemental Plates remain separate, but 
the references to placement into Lamarckian 
genera of taxa in the first part are not present, 
as they are so identified in the revised first part. 
In Hanley’s format there are four columns, the 
first being the figure number on the appropri- 
ate plate (determined by a header for each 
genus), followed by columns for “Wood's Lin- 
nean names”, “Lamarckian Genus,” and “Au- 
thority — Synonyms — Locality.” 

The real source of the confusion alluded to 
is the fact that new species names are first 
introduced in Linnaean (rendered as Linnean 
by Wood) genera and then seemingly reintro- 
duced in Lamarckian genera. Although it has 
sometimes been considered that the Lama- 
rckian genus was the genus in which a Wood 
name was first proposed, that position is un- 
dermined by the fact that Wood (1828b: [iii]) 
stated that the “plan pursued in the Index 
Testaceologicus [i.e., Wood 1828a] has been 
continued for the sake of uniformity.” It is no- 
table that in the “additional list” the “Lamarck- 
lan names” are cross-referenced to the 
Linnaean genera but he did not cross-refer- 
ence the Linnaean list. 

There are surprising few typographical er- 
rors in any of the editions. Only one has been 
noted involving a name referenced herein. 


APPENDIX C 


Notes on the Versions of the 2" Edition of 
Cuvier’s Le Règne Animal 


In our Introduction, the primary publications 
involved in this study were mentioned and 
briefly described. These works, as well as 
other peripheral publications, are here further 
listed, collated, and annotated. A complete bib- 
liography of Cuvier, including the many trans- 
lations and permutations of his Régne Animal, 
was compiled by Smith (1993). Her reference 
numbers are cited herein. 

In 1816 Georges Cuvier produced the first 
edition of Le Régne Animal, his classification 
of animals into four embranchments — verte- 
brates, articulates, mollusks and radiates. It 
was published in four volumes with only 15 
plates. The third volume, containing the crus- 
taceans, arachnids and insects, was written 
by P. A. Latreille. An enlarged second edition 
was begun in 1829. 


(1) Original Second Edition by Cuvier 


CUVIER, BARON [С. L. C. F. D.], 1829-1830, 
Le régne animal distribué d’apres son 
organisation, pour servir de base a l’histoire 
naturelle des animaux et d’introduction a 
l'anatomie comparée. Nouvelle édition, revue 
et augmentée. Paris: Déterville, 5 volumes 
[volumes 4 & 5 by P. À. Latreille]. 

1: xxxvili + 584 pp. (1829). 

2: xv + 406 pp. (1829). 

3: xvi + 504 pp., 20 pls. (1830). 

4: xxvii + 584 pp. [by P. A. Latreille]. Crustacés, 
arachnides et partie des insectes. (1829). 

5: xxiv + 556 pp. [by P. A. Latreille]. Suite et 
fin des insects (1829). 


Notes 


In the introduction to Vol. 3, Cuvier stated that 
in order to leave together all the portion of this 
work which Latreille agreed to undertake, he 
combined in this volume the mollusks and zoo- 
phytes in the order of his method. Volume 3 
contains an extensive “table alphabétique des 
auteurs cites dans cet ouvrage” (рр. 329—428), 
plate explanations (pp. 429-440), and an in- 
dex to all five volumes (pp. 441-504). This sec- 
ond edition is number 748 in Smith (1993: 185). 

The major criticism of Cuvier’s first edition 
was its paucity of illustrations. Even before this 


258 PETIT & COAN 


new edition was started, Edouard Guérin (b. 
1799 —d. 1874), who confusingly changed his 
name in 1836 to Guerin-Meneville, offered to 
produce an Atlas to offset this criticism. Ac- 
cording to Cowan (1971: 18), the original plan 
was to illustrate at least one species of every 
genus mentioned by Cuvier and Latreille. This 
work is discussed below. 


(2) The Griffith & Pidgeon Translation 


GRIFFITH, Е. & Е. PIDGEON, [1833]-1834, 
The Mollusca and Radiata. Vol. 12, In: Е. 
GRIFFITH, [1824]-1835, The Animal King- 
dom arranged in conformity with its organi- 
zation, by the Baron Cuvier, member of the 
Institute of France, &c. &c. &c. with supple- 
mentary additions to each order, by Edward 
Griffith, F. L. S., А. S., corresponding mem- 
ber of the Academy of Natural Sciences of 
Philadelphia, &c. and others. London: Whit- 
taker and Co., viii + 601 pp., 61 pls. [Frontis- 
piece (PI. 41 dated 1834 and not in List of 
Plates); Title-page; half-title-page; vii—vii (List 
of Plates); 1-138 (Mollusca, after Cuvier); 
139-434 (Supplementary Treatise on the 
Mollusca); 435-522 (The Zoophytes, or 
Animalia Radiata, after Cuvier); 523-594 
(Supplement to the Radiata); 595-601 (Al- 
phabetical List of the Figures of Mollusca); 
Mollusca Plates 1-41; Zoophytes Plates 1- 
20]. 


Notes 


This is Volume 12 in a series of 16 translat- 
ing and adding to Cuvier’s work. Details about 
the other volumes are given by Cowan (1969). 
The plates are not in numerical order but are 
placed in various positions within the work, the 
locations being shown on the List of Plates. 
The dates on the plates and what little is known 
about the dating of the work itself is discussed 
in our Introduction. 

Although there is an “Alphabetical List of the 
Figures of Mollusca,” there is no such list for 
the figures of Zoophytes. 

There are several oddities in the printing. 
According to McKerrow (1927: 81-82), press 
numbers were not used after 1823, but they 
appear to be present in this volume. Within 
each signature, with the exception of Ff, there 
is asmall number, 1, 10, or 13, below the text 
in various positions but always below the bot- 
tom line of text. Position within the signature 
varies, but there is only one number in each 
signature. 


This work was published in three forms: (1) 
Octavo, trimmed page size 13 x 21 cm. on copy 
at hand, with plain plates; (2) in the same size 
but with colored plates; (3) and in quarto, 
trimmed page size 16 x 24.8 cm, with colored 
plates. The terms octavo and quarto used by 
the publisher refer to the size, not manner of 
printing. The colored sets are only partially so, 
the plates illustrating sections of shells and 
terminology being uncolored. 

As discussed in our Introduction, some of the 
plates were copied from Guérin-Méneville and 
Blainville, with only 20 being original. Cowan 
(1969: 38) stated that Westwood (James 
Obadiah Westwood, b. 1805 — d. 1893) “cop- 
led many of Guérin's plates as well as supply- 
ing original ones”. It could thus be assumed 
that Westwood copied, or drew, all of the Mol- 
lusca plates, but there is at least one excep- 
tion. Mollusca Plate 14 is inscribed “W. Hawkins 
del”. This would be the noted natural history 
artist and sculptor Benjamin Waterhouse 
Hawkins (b. 1807 — d. 18897). Nissen (1969: 
108) did not attribute any other of the plates, 
but noted that this one was by Hawkins. 

~Westwood’s copies are quite acceptable, but 

some figures are less esthetic than Guérin- 
Meneville’s originals in that they have been 
reduced in size. In several instances, the fig- 
ures on two Guérin-Méneville plates were re- 
duced to fit onto one plate, or some of the 
multiple views of the same species were elimi- 
nated. Guerin-Meneville’s Mollusque plates 20 
and 21 contain 16 figures representing 14 spe- 
cies. Ten of these species were copied as 
Griffith & Pidgeon’s plate 33, with one figure 
per species, the omitted taxa being those un- 
colored in the Guérin-Méneville originals. 

Cuvier utilized extensive footnotes, and his 
format was copied in the Henderson edition 
(number 6 below). Griffith & Pidgeon, however, 
incorporated shortened versions of the foot- 
notes into the main text’s discussions of gen- 
era. They also added a large “Supplementary 
Treatise on the Mollusca” that occupies pages 
139-594 of their work. The author is unstated. 
The Supplement begins with a learned dis- 
course on the history of the development of 
the arrangement of Mollusca and continues 
with remarks on various genera, including 
ecology and physiology. An extensive glossary 
of descriptive terms is imbedded in the dis- 
cussions and many are illustrated. This 
Supplement is short on systematics; no new 
taxa are proposed, and surprisingly none of 
the new names introduced on the plates and 
in the Index are mentioned. 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 259 


The translation of Cuvier in the Henderson 
edition is remarkably similar to that of Griffith 
& Pidgeon, evidently due to the fact that both 
works translate the sentences with the same 
structure and in the same order as the origi- 
nal. Entire paragraphs are exactly alike, but 
occasionally a major difference will be found. 
Cuvier remarked on the estuarine habitat of 
potamides by stating that they were to be found 
in rivers “ou au moins à leur embouchure’. 
McMurtrie, translator of the Henderson and 
other editions, correctly translated this phrase 
as “or at least, their mouths.” Griffith & Pidgeon 
mistakenly translated it as “or at least their 
mantles’! Other similar differences are to be 
found, most often in references to morphol- 
оду. 

Although, as shown by Gruber (2004), this 
edition of Cuvier was Griffith's responsibility, 
the financial burden fell on the publisher, 
George Byrom Whittaker (b. 1793 — 4. 1847). 
Topham (2004) discussed Whittaker’s finan- 
cial affairs, stating that he “was even able to 
persist with the publication of his splendid edi- 
tion of Georges Cuvier’s Animal Kingdom (16 
vols., 1827—1835) for which the translation and 
the engraving of the plates alone cost £7000." 
That was an enormous sum at the time. Con- 
sider that in 1849 С. В. Sowerby II was being 
paid £1 per plate for the lithographs in Reeve’s 
Conchologia Iconica (Petit, 2007). 

Cowan (1969: 137) commented that the en- 
tire series was “meticulously translated, prob- 
ably mainly by Pidgeon.” 


(3) The 1988 Indian Reprint of Griffith & Pidgeon 


GRIFFITH, Е. & Е. PIDGEON, [1833]--1834 
[1988], Handbook of Mollusca and Radiata 
arranged by the Baron Cuvier with supplemen- 
tary additions to each order. Delhi, India: 
Biotech Books, iv + 602 pp., 60 pls. [reprint of 
the original Volume 12 with a new title page]. 


Notes 


In 1988, Volume 12 was reprinted by Biotech 
Books, Delhi. Neither the title page nor half- 
title page were reprinted. These were replaced 
by a new title page imprinted “Handbook of 
Mollusca and Radiata arranged by the Baron 
Cuvier with supplementary additions to each 
order by Edward Griffith and Edward Pidgeon.” 
The societal affiliations of the authors were 
omitted. A Contents page, not present in the 
Original, has been placed between the title 
page and the List of Plates. The reprint is cloth 


bound with a dust jacket. The dust jacket gives 
pagination as iv + 602 pp., 60 pls. The count 
of “602 pages” includes the blank verso of page 
601. Mollusca Plate 41, placed in the front of 
the original issue, has been moved opposite 
page 74. The reprinter’s count of 60 plates is 
unfortunately correct, as Plate 29 has been 
omitted. Also omitted on Page 601 is “End of 
Vol. XII, the printer's name and address, and 
signature letters”. The printing of the plates is 
very dark and unsatisfactory. 


(4) The Guérin-Méneville Illustrations 


GUERIN-MENEVILLE, F. E., 1829-1844, 
Iconographie de règne animal de С. Cuvier, 
ou représentation d’aprés nature de l’une des 
espéces les plus remarquables et souvent 
non encore figurees, de chaque genre 
d’animaux. Paris: J. В. Baillière, 3 Volumes 
(2 of plates, 1 of text): 


1. Planches des Animaux vertébrés. Half-t.p., 
t.p., ftsp. (portrait of Cuvier). 
Mammifères, 52 plates (A, B, 1-48, 11 bis, 
44 bis) 
Oiseaux, 70 plates (1-67, 22 bis, 23 bis, 36 
bis) 
Reptiles, 30 plates (1-30) 
Poissons, 70 plates (1-70; 31 misnumbered 
as 19) 
2. Planches des Animaux invertébrés. Half-t.p., 
t.p., ftsp. (portrait of Latreille). 
Mollusques, 38 plates (1-38) 
Annelides, 11 plates (1-10, 4 bis) 
Crustaces, 36 plates (1-35, 8 bis) 
Arachnides, 6 plates (1-6) 
Insectes, 110 plates (1-104, 24 bis, 25 bis, 
28 bis, 39 bis, 49 bis, 84 bis) 
Zoophytes, 25 plates (1-25) 
3. Texte explicatif. Half-t.p., t.p., 1-4 (Avis), 
[v] (Memorial page to Cuvier, Latreille and 
Delessert), [vii]xvi (Avant-propos). [The text, 
which is expanded plate explanations, is in 
sections, each preceded by a half-t.p. and a 
full t-p. for the group. These title pages, al- 
though unnumbered, are included in the num- 
bering in each section (i.e., actual text starts 
on р. 5 of each section)]: 
Mammifères, 36 pp. 
Oiseaux, 40 pp. 
Reptiles, 24 pp. 
Poissons, 44 pp. 
Mollusques, 29 pp. 
Annélides, 14 pp. 
Crustacés, 48 pp. 
Arachnides, 20 pp. 
Insectes, 576 pp. 


260 PETIT & COAN 


Notes 


This history of this work was discussed by 
Cowan (1971), who tried, with limited success, 
to date the various parts. Unfortunately for 
malacology, Cowan was primarily interested 
in the insects, which received his most detailed 
attention. The dates shown on the various title 
pages are not listed as they are, according to 
Cowan, meaningless. The plates, which ap- 
peared in advance of the text, bear legends 
from which many species of insects date. Al- 
though many of the plates of other Phyla are 
dated, the only mollusk plate bearing a date is 
Plate 1, which is dated 1829. Morrison (1971: 
566) dated Mollusques Plate 30 as 1832, but 
he did not give a collation or any reason for 
doing so. However, Guerin-Meneville’s 
Mollusques Plate 6 was cited by Deshayes 
(1832: 404), showing that at least some, if not 
all, of the mollusk plates were issued by that 
time. 

Cowan (1971) dated the plates by livraisons 
from 1829 to 1837, but there is no indication 
of what plates were issued in which livraison. 
He stated that the text was complete on Sep- 
tember 7, 1844. 

The exceptionally fine Mollusca plates in this 
work were drawn by Guerin-Meneville either 
by himself or in collaboration with Travies. One 
plate is stated to have been drawn “apres 
d’Orbigny”. The Mollusca plates were all en- 
graved by Giraud, except for one by “Lebrun 
et Giraud” and one by Pedretti. Not only are 
the plates finely drawn and engraved, but the 
coloring is remarkable. 

Advance copies were made available to 
Westwood for copying, as we note in the dis- 
cussion of Griffith & Pidgeon. The plates were 
also copied in the Henderson edition listed 
below. 

At the end of each section of explanatory text 
there is an alphabetical index. These indices 
have not been identified as such in the colla- 
tion as they are included in the pagination. 

There are 448 plates of animals. The figure 
450 listed by Cowan (1971: 29) obviously in- 
cludes the two portraits, as he gives the cor- 
rect number of plates for each section. 

Cowan listed three arrangements of the 
Iconographie that were deliberately planned 
and for which alternative title-pages were 
made available: 


Official (as the Atlas for the Règne Animal) 
Vol. 1. Planches des Animaux Vertébres. 
Frontispiece + 222 pls. 


Vol. 2. Planches des Animaux Invertébrés. 
Frontispiece + 226 pls. 
Vol. 3. Texte Explicatif. Approximately 930 pp. 


Simple (as a work in its own right; vol. 3 being 
very frequent) 
Vol. 1. Vertebrates. ftsp. + 144 pp., 222 pls. 
Vol. 2. Invertebrates (less Ins.). 176 pp., 116 
pls. 
Vol. 3. Insects. ftsp. + 576 pp., 110 pls. 


Specialist 

10 Volumes: a separate one for each Class; 
sizes varying from 24 pp., 11 pls. (Reptiles) to 
576 pp., 111 pls. (Insects). 

The molluscan part of this work (see below 
as “2 ed.”) was reprinted in Paris by Bailliere 
et fils in 1868: 74 pp., 36 pls. 


As mentioned above, Guérin added “- 
Méneville” to his name in 1836; here to avoid 
confusion, we use that combination surname 
throughout. There are some names on Griffith 
& Pidgeon plate legends that can cause con- 
fusion and which originated in the first edition 
of Guérin-Méneville. One such name appears 
on Griffith & Pidgeon plate 38, a plate com- 
posed of reduced figures from several Guérin- 
Méneville plates. This species is: 

Donax hilairea Guérin-Méneville, 1832: pl. 
30, fig. 4 — in legend as “Val. Col. Mus.” 
[probably from a museum label; never pub- 
lished by Valenciennes]. 

Donax hilairea — Griffith & Pidgeon, 1833: pl. 
38, fig. 9 [no author shown on plate legend]. 

Donax hilairea Valenc. — Griffith 8 Pidgeon, 
1834: 597. 

Donax denticulata Linn. — Guérin-Méneville, 
1844: 47 [non Donax denticulata Linnaeus, 
1758-6681: 

Donax hanleyana Philippi, 1847b: 84-85. 

Donax elongata Lamarck — [Deshayes in 
Guérin-Méneville & Deshayes], 1868: 59, 
pl. 29, fig. 4 [non Donax elongata Lamarck, 
18185501 

Donax hilairea Guerin-Meneville — Morrison, 
19712306; pl. 1 Ag 

This species was treated, as Donax hilairea 
Guérin-Méneville, by Morrison in a monograph 
on the western Atlantic Donax. The name 
Donax hilairea Guérin-Méneville, 1832, was 
suppressed in favor of the unfigured Donax 
hanleyana Philippi, 1847, in ICZN Opinion 
1372 (1986) based on an application by Narchi 
(1983). (The application was submitted by 
Narchi in 1975 and not published or acted 
upon by the Commission until after Morrison’s 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 261 


death in 1983 — see also Narchi, 1986). 

Also on Guérin-Méneville (1832) plate 30, fig. 
2, 2a, there is a figure of /socardia dussumierii 
Val. Col. Mus.” Not reproduced in Griffith & 
Pidgeon, it is mentioned here as an example 
of what may be many Guérin-Méneville names 
that have escaped notice. Neither Donax 
hilairea nor Isocardia dussumierii were listed 
by Sherborn. In the 1844 Guérin-Méneville 
Texte, this figure is listed as a form of /. 
moltkiana (Gmelin, 1791: 3303-3304). With- 
out mentioning the name used on the plate, it 
is stated that the name thereon was not 
adopted by Valenciennes as it is of no scien- 
tific value. The figure to which /socardia 
dussumierii applied was even later identified 
as Isocardia vulgaris Reeve, 1845 by 
Deshayes in [Guérin-Méneville & Deshayes] 
(1868: 59, pl. 29, fig. 2a, b). It is obvious that if 
they are conspecific, Reeve's name would be 
junior to /. dussumierii. The eponym dussumieri 
was used by Cuvier & Valenciennes for a num- 
ber of species of fish, but this usage for the 
mollusk was never published by Valenciennes. 
The resolution of the status of such Guérin- 
Méneville names is beyond the scope of this 
paper. 


(5) Guérin-Méneville “2"* edit.” 


[GUÉRIN-MÉNEVILLE, G. F. E. & G. P. DES- 
HAYES], 1868, Les mollusques décrits et 
figurés d’après la classification de Georges 
Cuvier mise au courant des progrés de la 
science. XXXVI planches représantant en 
520 figures Dessinées d’aprés nature et 
gravées sur cuivre Les espèces les plus 
remarquables de ces animaux avec un texte 
descriptif. J. В. Paris: Bailliére et fils, 74 pp., 
36 plates. 


Notes 


This rare work, basically the mollusk plates 
of Guerin-Meneville’s /conographie, does not 
identify the author(s) on the title page or else- 
where. Copies have been located in four li- 
braries: Smithsonian Institution, the personal 
library of Scott Jordan, the University of 
Glasgow Library, and the Bibliotheque National 
in Paris. In the libraries, it is catalogued under 
Cuvier, but catalogues of the latter two show 
that the text is by G. P. Deshayes. Smith (1993: 
206, item 803) listed the author as Guérin- 
Meneville, with his name in brackets, and in a 
footnote referenced the Bibliotheque National 


catalogue notation “Refonte partielle de 
Plconographie.” Cuvier certainly did not write 
the annotated plate explanations, the only text 
involved. We consider that authorship of the 
entire work should be attributed to Guérin- 
Méneville and Deshayes as shown in our 
header (in square brackets as their names are 
not shown on the work) and that the text should 
be attributed to Deshayes. The fact that 
Guérin-Méneville and Deshayes drew the fig- 
ures for one of the new plates in this work 
shows that both contributed to its production. 

Cowan (1971: 22) mentioned a second edi- 
tion having been frequently referred to by 
Guerin-Meneville, but that no such edition had 
been found although: 

“Guerin himself frequently referred to figures 
‘wrongly named in the first issue of plates, 
which have been corrected in our second edi- 
tion’ [here Cowan lists the insect plates af- 
fected]. No catalogue listing a second edition 
has been found. No copies of revised plates 
have been seen. It is possible that nomencla- 
tural problems may occur. All references seen 
are to plates dated before July 1835 or un- 
dated.” 

No source was given for the quote from 
Guérin-Méneville, and no complete second 
edition has been located. It is not known if this 
volume is indeed part of a “second edition” but 
it is logical to refer to it as such. 

This work came to our attention because 
Morrison (1971) cited a name and figure from 
“Guerin, 1868, Icon. du Regne Animl., reprint.” 
The copy used and cited by Morrison is in 
Smithsonian Institution, specifically in the 
mollusk library of the National Museum of 
Natural History. 

Smith mentioned that in some catalogues the 
work appears under Cuvier as author and that 
the explanation of plates is a different text from 
the /conographie. A detailed comparison of 
these “editions” was beyond the scope of 
Smith's work, and it is not surprising that she 
made small errors about their differences. She 
Stated that “with exception of pl. 2, plates are 
same as those in the Iconographie, although 
[2"“ edition] lacks last plates of rec. 802 [= 1° 
edition] in the copy seen.” While the 1* edition 
has 38 mollusk plates, the 2™ edition contains 
only 36. Smith was correct about Plate 2 be- 
ing different, but she could not account for the 
absence of the last two plates, nor did she note 
any other differences. 

Four plates of the 1* edition — plates 2, 3, 
37, 38 — were not reprinted in the 2™ edition. 


262 PETIT & COAN 


The plates in the 2" edition can be accounted 
for as: 

Plate 1 - Same as in 1* edition 

Plate 2 —A new plate figuring living and fos- 

sil cephalopods, drawn by Thiolat and 

Lackerbauer. 

Plates 3-35 — These are plates 4—36 of the 

15 edition, each being renumbered, with the 

result that the 2"* edition versions have a 

plate number one digit lower. 

Plate 36 — A new plate of Dentaliidae, drawn 

by Guérin-Méneville & Deshayes. This Class 

was omitted from the 1° edition. 

The plates for the 2" edition were not 
reengraved. The figures are within a single line 
border with book title and plate number above 
the top border and the legend with species 
names below the bottom border. Also, imme- 
diately below the bottom border line appear 
the names of the artist(s), printer and engraver. 
With the exception of the artisans’ names and 
the plate numbers, all matter outside of the 
border was deleted for the 2" edition. Of 
course, all of the plate numbers, except for 
Plate 1, had to be changed. There is no title 
above the border as in the 1* edition. The 
name of the publisher, J. В. Balliére et fils, a 
Paris, appears in the space where the legend 
was originally. In the 2" edition, the names of 
the figures appear only in the plate explana- 
tions, the only text in the volume. 

Due to the long delay between the printing 
of the plates and the text (i.e., the plate expla- 
nations) of the first edition, the names in the 
text do not always match the plate legends. In 
the 2° edition, many names have been 
changed again from the 1* edition. As an ex- 
ample, we note above, all for the same figure: 

Donax hilairea — 1* edition, plate legend 

Donax denticulata — 1* edition, text 

Donax elongata — 2" edition, text 


(6) The Henderson Edition 


CUVIER, BARON [6. L. C. F. D.], [1833]-1837, 
The Animal Kingdom, arranged according to 
its organization, serving as a foundation for 
the natural history of animals, and as an in- 
troduction to comparative anatomy. The 
Crustacea, arachnides, & Insecta, by M. 
Latrielle, ... translated from the latest French 
Edition. London: G. Henderson, 4 volumes 
(in 8). 


Text 
1. [1833]-34. Mammalia — birds. Frontispiece 
(portrait of Cuvier), pp. i-ii (advertisement; 


= Preface), + i-xlvii [i-xvi (Memoir of Cuvier); 
xvii-xxvill (Preface to 1* edition); xxix—xxxi 
(Preface to 2" edition); xxxiii-xlviii (index, 
corrigenda)] + 1-380. 

2. [1833]-34. Reptiles — Fishes. Frontispiece 
(portrait of Latreille), pp. xxxv + 412 [pp. 1- 
64 — published 1833]. 

3. 1834. Mollusca — annelides — Crustacea — 
arachnides and Insecta. Frontispiece (por- 
trait of Buffon), pp. xxxv + 472. 

4. 1836. Insecta — zoophytes. Frontispiece 
(portrait of Audubon), pp. 1-й (memorial to P. 
A. Latreille, who had died in 1833) + i-xliv + 
544. 


Plates 

1. 1837. Mammalia — birds. xviii pp. (plate ex- 
planations). Mammalia, 80 pls. [numbered 
1-63 + 4*, 5*, 6*, 6 bis, 7*, 2% pl. 24, 30 bis, 
2™ pl. 30 bis, 46 bis, 47 bis, 48 bis, 49 bis, 
50 bis, 53 bis, 54 bis, 54 ter, 57 bis]; Avis, 
117 pls. [numbered 1-83, 9899, + 2 bis, 3 
bis, 4 bis, 6 bis, 2" pl. 18, 2° pl. 24, 2" pl. 
212 Mb18,:29 biset3tibisy3S bisy 35er, 27 
pl. 42, 42 bis, 2% pl. 44, 2% pl. 45, 3 pl. 45, 
46 bis, 46 ter, 47 bis, 50 bis, 56 bis, 2" pl. 
61, 61 bis, 66 bis, 68 bis, 69 bis, 2"* pl. 69 
bis, 69 ter, 70 bis, 70 ter, 71 bis]. 

2. 1837. Reptiles — fishes. Frontispiece (por- 
trait of Linnaeus), xxii pp. (plate explana- 
tions). Reptilia, 43 pls. [numbered 1-40 + 
2™ pl. 6, 25 bis, 26 bis]; Pisces, 154 pls. 
[numbered 1-9, 11-20, 2336, 38—40, 42-80, 
+ 8 bis, 10 bis, 12 bis, 13 bis, 14 bis, 16 bis, 
17 ter, 18 bis, 2% pl. 18 bis, 18 ter, 19 bis, 20 
bis, 21 bis, 22 bis, 23 bis, 24 bis, 24 ter, 2" 
pl. 25, 25 bis, 25 ter, 2" pl. 26, 26 bis, 26 ter, 
26 quar, 27 ter, 2" pl. 27 ter, 27 quar, 28 bis, 
2° 91:28:bisı3"" pl. 28 bis, 2ВЧег 20% р 28 
ter, 28 quar, 2" pl. 28 quar, 2" pl. 29, 29 bis, 
29 ter, 2" pl. 29 ter, 30 bis, 2" pl. 30 bis, 30 
ter, 2" pl. 30 ter, 30 quar, 31 bis, 31 ter, 31 
quar, 32 bis, 32 ter, 2"? pl. 32 ter, 32 quar, 2" 
pl. 33, 33 bis, 33 ter, 33 quar, 34 bis, 34 ter, 
34 quar, 35 bis, 35 ter, 35 quar, 36 ter, 37 
ter, 37 quar, 38 bis, 38 ter, 38 quar, 39 ter, 
40 bis, 41 bis, 42 bis, 42 ter]; Pisces Osteol- 
ogy Plates, 1-8 [double fold-out plates]. 

3. 1837. Mollusca — annelides — Crustacea and 
arachnides. Frontispiece (portrait of William 
Kirby), xxiv pp. (plate explanations). Mol- 
lusca, 79 pls. [numbered 1-44, 2 bis, 4 bis, 
4 ter, 16 bis, 16 ter, 22 bis, 22 ter, 23 bis, 23 
ter, 23 quar, 24 bis, 2" pl. 24 bis, 24 ter, 25 
bis, 25 ter, 26 bis, 26 ter, 31 bis, 2° pl. 32, 
32 bis, оном ter, 21 
pl. 32 ter, 2" pl. 33, 33 bis, 34 bis, 35 bis, 36 


MOLLUSCAN TAXA IN GRIFFITH & PIDGEON (1833-1834) 263 


bis, 37=bis,374ter, 27, p 1243,43 He 2" pl, 
44, 44 ter]; Annelides, 11 pls. [numbered 110, 
4 bis]; Crustacea, 59 pls. [numbered 1-42, 
24 bis, 25 bis, 27 bis, 28 bis, 29 bis, 31 bis, 
31 ter, 2% pl. 31 ter, 32 bis, 32 ter, 33 bis, 33 
ter, 34 bis, 35 bis, 35 ter, 36 bis, 39 bis]; 
Arachnides, 51 pls. [numbered 1-29, 1 bis, 
2 bis, 3 bis, 5 bis, 6 bis, 7 bis, 8 bis, 9 bis, 9 
ter, 10 bis, 10 ter, 11 bis, 11 ter, 12 bis, 13 
bis, 16 bis, 16 ter, 17 bis, 17 ter, 18 bis, 18 
ter, 24 bis]. 

4. 1837. Insecta — Zoophytes. Frontispiece 
(portrtait of William Spence), xxxii pp. (plate 
explanations). Insecta, 120 pls. [numbered 
1-115, 35 bis, 38 bis, 59 bis, 2" pl. 94, 98 
bis]; Zoophytes, pls. 1-24. 


Notes 


This “Henderson Edition” is so referenced 
in the on-line catalogue of The Natural His- 
tory Museum, London, obviously for the name 
of the publisher. For convenience, it is so re- 
ferred to herein. All volumes were published 
by G. Henderson, except Text Volume 4, which 
was published by E. Henderson. The firm, 
under both names, was at the same Ludgate 
Hill, London, address. The firm is not listed, 
under either name, by P. A. H. Brown (1982). 
This is Smith’s number 756 (Smith, 1993: 189). 

This is an English translation of the 2" 
French edition prepared by the American Dr. 
Henry McMurtrie (published as M’Murtrie; b. 
1793 — d. 1865). The text of this edition is, 
according to Smith, identical to an edition pub- 
lished in Philadelphia in 1831 (Smith, 1993: 
188, number 753). As a sign of the times, 
McMurtrie also produced a one-volume bowd- 
lerized edition in 1832 (Smith, 1993: 188-189, 
number 754), in which he stated that: “The 
whole has now been sedulously, ... and so 
thoroughly, expurgated that it may be placed 
in the hands of females, without the slightest 
fear of their encountering a word or idea that 
could offend the most fastidious delicacy ...” 
(McMurtrie, 1832: vi). 

Text volumes are imprinted 1834. Woodward 
(1903: 410) stated that “the text is dated 1834, 
but Vol. | & Il, pp. 1-64, appeared in 1833.” 
Evidence for this is not stated, and it is not 
evident in the work. In fact, in Volume |, pages 
1-48 are on different paper than the rest of 
the volume. Woodward also stated that “the 
four volumes of plates are dated 1837, the last 
figure having apparently been altered”. Under 
magnification it can be seen that the right por- 


tion of the “leg” of the 7 may have been part of 
a 4 but if altered it is a superb job of partial 
reengraving. Volume 4 is dated 1836. Fortu- 
nately, no new names are believed to date from 
this edition, so the precise date is of little im- 
port. This work is number 756 in Smith (1993: 
189-190), who did not list the two-page me- 
morial to Latrielle in Volume 4. 

As can be inferred from the weird number- 
ing, counting the plates in this set is difficult. 
Nissen (1969: 108) errs in listing 156 instead 
of 154 Pisces plates, as allowance was not 
made for the fact that there are no plates 10 
or 27, only “supernumerary” plates with those 
numbers. Aves are listed as having 114 plates 
instead of 117 as “2" Plates” 18, 42 and 44 
were not included in his count. Arachnides 
were listed as having 99 plates instead of 51. 
This error was obviously typographical as the 
plates are listed by number and can be 
counted. The Insecta are listed as having 124 
plates instead of 120, evidently because the 
plate explanations list Plates 32 and 72 as 
each being three plates, so listed as these two 
plates are line drawing of parts of insects, each 
plate divided into thirds with the figures in each 
third separately numbered. The total number 
of plates in the entire work is 738. 

Smith (1993: 189) also erred in listing Plate 
Volume 2 as containing 201 plates instead of 
205 and Plate Volume 4 to have 150 plates 
instead of 144. This latter number may be par- 
tially explained by two plates being divided into 
three numbered parts listed separately in the 
Explanations, as mentioned in the previous 
paragraph. Smith also stated that the “num- 
ber of plates varies among copies seen.” This 
does not make sense as each of the plate 
volumes has a detailed Table of the Plates list- 
ing each plate by number with the names ac- 
companying the figures thereon. 

The portrait of Cuvier in Volume 1, engraved 
on steel by Brown (identity unknown) is a copy 
of a portrait that appeared in Volume 1 of 
Guérin-Méneville drawn and engraved by 
Bertonnier (Pierre-François Bertonnier, b. 
1791 — d. unknown). Bertonnier is shown as 
artist on the Brown engraving. 

The plates of all groups have not been in- 
spected in detail, but those of Mollusca in this 
edition are very poor, both in detail and in color. 
The 38 plates of Mollusca from Guérin- 
Méneville were copied with varying degrees 
of success. They are not the copies made for 
Griffith & Pidgeon and are much inferior. Of 
the 38, 16 (17-26, 33-38) were reversed by 


264 PETIT & COAN 


the copier. It appears that the plates from 
Guérin-Méneville may have been received in 
groups as printed and that they were then 
placed in the hands of various copiers. This 
observation is based on the fact that the qual- 
ity of the copies differs from lot to lot, becom- 
ing progressively worse. A cursory examination 
shows that copied Guérin-Méneville plates 
were utilized throughout and that some plates 
in other groups were also reversed and poorly 
colored. 


There are signs of poor editing and proof 
reading in this work. A cursory review found 
such errors as Buccinum undulatum for B. 
undatum; B. tessellatus for tessulatus; B. 
erinaceous for erinaceus; Murex ricinis for M. 
ricinus, M. singulatus for cingulatus; and 
Terabra for Terebra; all on pp. 69-72. On the 
other hand, some improvements were made 
such as using the genus name Potamida, 
whereas Cuvier used only the vernacular 
“potamides”. 


MALACOLOGIA, 2008, 50(1-2): 265-277 


MOLLUSCAN NAMES AND MALACOLOGICAL CONTRIBUTIONS OF 
WOLFGANG KARL WEYRAUCH (1907-1970) WITH A BRIEF BIOGRAPHY 


André F. Barbosa’, Valdemar К. Delhey? & Eugene V. Coan™ 


ABSTRACT 


Wolfgang Karl Weyrauch (1907-1970) studied land and freshwater gastropods of South 
America, mainly taxa belonging to the families Camaenidae, Charopidae, Clausiliidae, 
Endodontidae, Helicinidae, “Hydrobiidae”, Orthalicidae, Pupillidae, Scolodontidae, 
Subulinidae, and Urocoptidae. Here we list the 198 molluscan names introduced by 
Weyrauch and all of his publications of malacological interest. A brief biography of Weyrauch 
and a list of taxa named for him are also provided. 

Key words: Mollusca, Gastropoda, Wolfgang Karl Weyrauch, bibliography. 


RESUMEN 


Wolfgang Karl Weyrauch (1907-1970) se 
dedico al estudio taxonömico de gasteröpodos 
sudamericanos terrestres y dulceacuicolos, en 
su mayoria integrantes de las familias 
Camaenidae, Charopidae, Clausiliidae, 
Endodontidae, Helicinidae, “Hydrobiidae”, 
Orthalicidae, Pupillidae, Scolodontidae, 
Subulinidae y Urocoptidae. Se presenta el 
inventario de los 198 nombres de moluscos 
introducidos por Weyrauch y una lista 
completa de sus trabajos de interés 
malacologico. Se incluye ademas una breve 
biografia de Weyrauch y los nombres de 
especies descriptas en su honor. 

Palabras clave: Mollusca, Gastropoda, 
Wolfgang Karl Weyrauch, bibliografia. 


INTRODUCTION 


Wolfgang Karl Weyrauch (1907-1970), one 
of the most proficient malacologists of his time, 
contributed substantially to our knowledge of 
South American mollusc diversity. This contri- 
bution is reflected by the large number of 
Weyrauch specimens deposited in museums 
worldwide. 

Weyrauch sometimes distributed named 
shells, some labeled as type specimens, be- 
fore formally publishing their descriptions. Un- 


fortunately, Weyrauch died before publishing 
many manuscript names, and according to Zilch 
(1970), Weyrauch intended to describe of 50 
new gastropod species just before his death in 
a manuscript that was never published. 

Shortly after Weyrauch’s death, Zilch (1970) 
published a bibliography of his entomological 
and malacological papers, with a list of his new 
mollusc names. However, this paper is in a 
journal available in few libraries in South 
America, where Weyrauch conducted his ma- 
lacological research, and it omits one 
Weyrauch paper and four taxa. It is not sur- 
prising that curators are often puzzled as to 
the validity of some labels and names applied 
to Weyrauch’s material. 

This paper inventories of the molluscan 
names made available by Weyrauch and lists 
his publications of malacological interest. We 
also provide a partial list of the type material of 
his species based on the original descriptions 
and on discussions with malacological cura- 
tors. A brief summary of Weyrauch’s life is pro- 
vided following Aguilar (1970), Duarte (1970), 
Lamas (1981), Willink (1999), and Zilch (1970). 


BIOGRAPHY 
Born on December 7, 1907, in Elberfeld, 


Germany, Wolfgang Karl Weyrauch obtained 
a PhD in 1929 at Friedrich Wilhelm Univer- 


‘Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renovaveis - IBAMA, Coordenacäo Geral de Petröleo e Gas, 
Praga XV de Novembro, 42, 9° andar. 20010-010 Rio de Janeiro, RJ, Brasil 
“Departamento de Biologia, Bioquímica y Farmacia, Universidad Nacional del Зиг. San Juan 670, (8000) Bahia Blanca, 


Argentina 


¿Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, California 93105-2936, U.S.A. 


“Corresponding author: gene.coan@sierraclub.org 


266 BARBOSA ET AL. 


sity, Berlin, completing a thesis on insect neu- 
rophysiology. From 1928 to 1929, he worked 
as Assistant in Zoology to Richard Hesse, one 
of the pioneers of animal ecology. Between 
1931 and 1943, Weyrauch worked for the Ger- 
man Council of Scientific Research doing en- 
tomological and ecological field studies in 
Europe and North Africa. By 1938, Weyrauch 
was in Peru, where he worked as entomolo- 
gist at the Estacion Agricola de La Molina in 
Lima until 1946, when he began work at the 
Estacion Experimental Agricola de Tingo Maria, 
also in Lima. During the Second World War, 
Weyrauch spent a few years in Texas, U.S.A., 
where he did field work in entomology and 
malacology. From 1948 to 1961, he worked at 
the Museo Nacional de Historia Natural, 
Facultad de Ciencias, Universidad Mayor de 
San Marcos, Lima, where he was a professor 
of systematic zoology, animal ecology, Zooge- 
ography, and genetics. From 1959 to 1961, he 
also worked as professor of agricultural Zool- 
ogy at the Facultad de Agronomia de la 
Pontificia Universidad Catolica del Lima. In 


FIG. 1. Wolfgang Karl Weyrauch, 1907-1970. 
Image from portrait in the Fundacion Miguel Lillo, 
Tucuman, Argentina; also reproduced in Aguilar 
(1970). 


1962, Weyrauch moved to Argentina, and be- 
came a researcher and professor at the 
Instituto Miguel Lillo, Tucumän, where he re- 
mained until his death of a heart attack on July 
2121970. 

Despite his initial dedication to entomology, 
Weyrauch changed the focus of his research 
to the Mollusca when he moved to South 
America, because this group better satisfied 
his interests in systematics, ecology, and bio- 
geography. He collected extensively, includ- 
ing marine molluscs, and sent duplicate 
material to museums and private collectors 
throughout the world. He discovered many 
new taxa, which he often studied in collabora- 
tion with other experienced malacologists and 
collectors, such as Joseph Bequaert, Fritz 
Haas, Miguel Klappenbach, Henry A. Pilsbry, 
Fritz Seidl, and Adolf Zilch. His work focused 
primarily on the terrestrial and freshwater gas- 
tropods of Argentina, Bolivia, Brazil, Ecuador, 
Peru, Uruguay, and Venezuela. 

Weyrauch’s first malacological contribution 
was the description of four new gastropod 
species and appeared in a paper by Zilch 
(1954). From 1956 until 1967, he published 
23 papers of malacological interest, mostly 
descriptions of new taxa based chiefly on shell 
characters. Papers with malacological content 
but without descriptions of new taxa are Meyer 
& Weyrauch (1965a, b), Weyrauch (1967c), 
and Weyrauch & Coronado (1958). 

According to Willink (1999), Weyrauch was 
not very kind and rejected the opinions of many 
colleagues and students. Aguilar (1970) de- 
scribed him as a very devoted and sage pro- 
fessor. An indefatigable worker, Weyrauch 
arrived in the laboratory at 3 or 4 p.m., and 
then worked all night, until his wife, Imelda 
Valdizan de Weyrauch, came to take him home 
in the morning. Acompulsive smoker, his labo- 
ratory was a noxious environment, contribut- 
ing to an antisocial way of life. 

In the Mollusca, Weyrauch described 119 
species, 45 subspecies, 10 genera, and 20 
subgenera (one unavailable), and he renamed 
four homonyms. As a result of his wide aca- 
demic collaboration, several species were 
named in his honor, including at least five mol- 
luscs, one scorpion, and a number of insects. 

Weyrauch’s private collection (WW) is now 
mainly in the Fundacion Miguel Lillo (FML), 
Argentina (Drahg & Cuezzo, 1999), and the 
Museo de Historia Natural (MHN), Peru. The 
Field Museum in Chicago, Illinois, U.S.A., pur- 
chased many lots from his widow after his 
death. The present status of the Angel E. Florez 
collection (Cuzco, Peru) is unknown to us. 


WOLFGANG KARL WEYRAUCH 267 


Collection Abbreviations 


AM See next entry 


AMS The Australian Museum, Sydney, 
Australia [originally indicated by 
Weyrauch as AM] 

ANSP Academy of Natural Sciences of 
Philadelphia, Pennsylvania, U.S.A. 

BML See next entry 

BMNH British Museum (Natural History) col- 
lection, The Natural History Museum, 
London, U.K. [originally indicated by 
Weyrauch as BML] 

CAS California Academy of Sciences [In- 
vertebrate Zoology type collection], 
San Francisco, California, U.S.A. 

CNHM See next entry 

FMNH Field Museum of Natural History, 


Chicago, Illinois, U.S.A. [originally 
indicated by Weyrauch as CNHM, 
Chicago Natural History Museum] 
FEL Weyrauch's abbreviation for the col- 
lection of Frederik Elisa Loosjes, 
Wageningen-Hoog, The Nether- 
lands; now in the Nationaal Natuur- 
historisch Museum [Naturalis], 
Leiden, The Netherlands 
Fundacion Miguel Lillo, Tucuman, 
Argentina [formerly Instituto Miguel 
Lillo, with lots referred to by Wey- 
rauch as IML] 
IML See preceeding entry 
MCZ Museum of Comparative Zoology, 
Harvard University, Cambridge, Mas- 
sachusetts, U.S.A. 
Museo de Historia Natural, Lima, Peru 
Museu Nacional, Universidade Fed- 
eral do Rio de Janeiro, Brazil 
Museu Rio-Grandense de Ciéncias 
Naturais, Porto Alegre, Brazil 
National Museum of Canada, 
Ontario, Canada 
Nationaal Natuurhistorisches Mu- 
seum, Leiden, The Netherlands 
[originally indicated by Weyrauch as 
RML, Rijks Museum van Natuurlijke 
Histoire] 
RML See previous entry 
SBMNH Santa Barbara Museum of Natural His- 
tory, Santa Barbara, California, U.S.A. 


FML 


MHN 
MNRJ 


MRCN 
NMC 


NNM 


SMF Naturmuseum Senckenberg, Frank- 
furt am Main, Germany 
USNM United States National Museum col- 


lection, National Museum of Natural 
History, Smithsonian Institution, 
Washington, D.C., U.S.A. 


WW Original Weyrauch collection numbers 
ZMB Museum für Naturkunde, Humboldt- 
Universitat Berlin, Germany 

Other Abbreviations 


HT Holotype 
RT Paratype 


MOLLUSCAN NAMES INTRODUCED BY 
W. К. WEYRAUCH 


Weyrauch’s new taxa — species, subspecies, 
genera, and subgenera — appear in alphabeti- 
cal order. The type material he mentioned or 
that has come to our attention for each species 
is given. The entries in bold face indicate data 
that is considered to be reliable, either because 
we have verified it with curators of the instutions 
or data provided by Zilch (1970) and Neubert 
& Janssen (2004) for the SMF, by Drahg & 
Cuezzo (1999) for the FML, and by Köhler 
(2007) for the ZMB. 

It can then be noted that some specimens in- 
dicated by Weyrauch to be in the FML, includ- 
ing some holotypes, or to have been sent to 
various other institutions have not yet been rec- 
ognized. However, we have also noted instances 
when the original numbers in Weyrauch’s col- 
lection differ from those incorporated into the 
FML collection to allow recognition of parts of 
these lots in other collections. Original Weyrauch 
numbers also indicate instances when the 
present localition of type material is unknown. 

In three cases here, species names are here 
given with umlauts, as in Weyrauch’s original; 
under the Code of Zoological Nomenclature 
(ICZN, 1999: Art. 32.5.2), such accents are to 
be eliminated in current usage. Some institu- 
tions have changed names since Weyrauch’s 
time; these changes are indicated in the ab- 
breviations listed above. 

The family allocations here follow our current 
undersanding of the placement of the genera 
he employed, translated into modern terminol- 
ogy, as given by Bouchet & Rocroi (2005). Ina 
few cases, recent literature indicates the cor- 
rect placement of some of his species. 

Because the descriptions of these taxa are 
accessible in reasonably common journals, we 
have not attempted to repeat the detailed type 
localities, which were often provided in lengthy 
German or Spanish sentences, with paratypes 
sometimes from multiple stations. Instead, only 
the country and region are given here. 


268 BARBOSA ET AL. 


acobambensis, Drymaeus — Weyrauch, 
1967b: 482-483, pl. 3, figs. 24, 25; central 
Peru [HT SMF 155694; PT FML 1524, 4140, 
4141, 4142, 10253]. Orthalicidae. 

aequistriata, Gracilinenia —\Weyrauch, 1956a: 
110-113, pl. 6, fig. 2; Valle de Chanchamayo, 
Peru [HT SMF 156390 (ex WW 1531); PT 
ANSP 319070, FEL, FMNH 54018, MCZ, 
SMF, Werner Blume collection (now in the 
SMF), ZMB 101651, WW]. Clausiliidae. 

Aeropictus [subgenus of Plekocheilus] — 
Weyrauch, 1967b: 465-467. Type species 
by original designation: Bulimus veranyi L. 
Pfeiffer, 1847. Orthalicidae. 

agitata, Pseudoglandina — Weyrauch, 1967b: 
486-487, pl. 4, fig. 53; central Peru [HT FML 
1066; PT FML 10238 (ex WW 1614, 10609), 
SMF 162138]. Amphibulimidae. 

agueroi, Bostryx (Peronaeus) — Weyrauch, 
1960c: 126, pl. 12, figs. 39-41; central Peru 
[HT SMF 162150; PT SMF 162151, 325576, 
FML 1462, FMNH 107836, 216889, 216890, 
USNM, WW 1268]. Orthalicidae. 

aguilari, Bostryx (Bostryx) — Weyrauch, 1967a: 
349-351, pl. 1, figs. 2-6; near Lima, Peru 
[HT SMF 162163; PT SMF 162164, 162165, 
162166, FML 1085, 1425, 4000, 4002, 4003, 
4004, 4005, FMNH 216900, 216901, 
216902, 216916]. Orthalicidae. 

albicolor, Peruinia — Weyrauch, 1957: 13-15, 
pl. 1, fig. 11; central Peru [HT SMF 155719; 
PT FMNH 217015, WW 1818]. Clausiliidae 

albocostata, Temesa (Temesa) — Weyrauch, 
1963b: 270-272, pl. 1, figs. 1, 2; central Peru 
[HT SMF 162108; PT SMF 162109, 162110, 
FML 1038, FMNH 107844, 217001, SBMNH 
83314, USNM, WW 3322]. Clausiliidae. 

altispira, Euglandina — Weyrauch, 1960а: 25— 
26, pl. 4, fig. 21; northern Peru [HT SMF 
162003; PT FML 3200, FMNH 216197]. 
Spiraxidae. 

altorum, Mesembrinus (Mormus) expansus — 
Weyrauch, 1958: 129, pl. 7, figs. 12, 13; cen- 
tral Peru [HT SMF 156295; PT SMF 156294, 
FML 1333]. Oleacinidae. 

altorum, Systrophia (Systrophia) — Weyrauch, 
1967a: 428—429, pl. 6, fig. 80; central Peru 
[HT FML 10668; PT FML 3962]. 
Systrophiidae. 

Andiniella [subgenus of Steeriana] — 
Weyrauch, 1958: 93-94. Type-species by 
original designation: Andinia (Ehrmanniella) 
flammulata Loosjes, 1957. Clausiliidae. 

andivagus, Naesiotus — Weyrauch, 1956b: 56, 
pl. 1, fig. 2; central Peru [HT FML 1461; PT 
ANSP 195000, SMF 155695]. Orthalicidae. 


angelmaldonadoi, Bostryx (Bostryx) modestus 
— Weyrauch, 1960а: 31-32, pl. 3, figs. 11, 
12; central Peru [HT SMF 155595; PT SMF 
162034, 162035, FML 12842 (ex WW 842)]. 
Orthalicidae. 

angiportus, Newboldius — Weyrauch, 1960b: 
53-55, pl. 8, figs. 5, 6; central Peru [HT SMF 
162045; PT SMF 161252, 162096]. 
Orthalicidae. 

angispira, Bostryx (Bostryx) obliquiportus — 
Weyrauch, 1960c: 123, pl. 11, fig. 9; central 
Peru [HT SMF 162183]. Orthalicidae. 

angustus, Bulimulus (Bulimulus) vesicalis — 
Weyrauch, 1966: 45-46, fig. 5; Rio Grande 
do Sul, southeastern Brazil [HT MRCN 
1576a; PT MRCN 1576, FML 10741]. 
Orthalicidae. 

araozi, Bulimulus (Bulimulus) — Weyrauch, 
1956c: 149-150, pl. 11, fig. 8; central Peru 
[HT SMF 155304; PT FML 2131]. Ortha- 
licidae. 

argentinus, Pupoides (Ischnopupides) 
chordatus — Weyrauch, 1964b: 37-38, fig. 
1; Jujuy, Argentina [HT FML 522a; PT FML 
522]. Pupillidae. 

Bakerilymnaea —Weyrauch, 1964a: 169. nom. 
nov. pro Nasonia F. C. Baker, 1928, non 
Ashmead, 1904. Lymnaeidae. 

bambamarcaénsis, Naesiotus (Naesiotus) — 
Weyrauch, 1960a: 37-38, pl. 6, fig. 38; north- 
ern Peru [HT SMF 156220; PT FML 3075, 
CAS 64085, USNM]. Orthalicidae. 

basiplanata, Epiphragmophora — Weyrach, 
1960a: 43-44, pl. 6, fig. 39; east-central Peru 
[HT SMF 162033]. Epiphragmophoridae. 

beltrani, Bostryx (Peronaeus) адиего! — 
Weyrauch, 1964b: 52-53, figs. 9, 10; cen- 
tral Peru [НТ SMF 162152; PT SMF 162153, 
FML 1084, FMNH 107849, 216891, 216892, 
USNM, WW 3116]. Orthalicidae. 

bequaerti, Drymaeus — Weyrauch, 1956c: 
154-155, pl. 11, figs. 12-14; central Peru [HT 
SMF 155309; PT SMF 155305, 155310, FML 
511, FMNH 30871]. Orthalicidae. 

bequaerti, Temesa (Neniatracta) — Weyrauch, 
1957: 25-27, pl. 1, fig. 12; northern Peru [HT 
SMF 155720; PT SMF 155721, ANSP 
204507, FEL, ЕММН, МСА 211952, USNM, 
WW 2010, 2010A, unspecified “private col- 
lections in U.S.A.”]. Clausiliidae. 

Bequaertinenia — Weyrauch 1964c: 150. Type 
species by original designation: Temesa 
(Neniatracta) bequaerti Weyrauch, 1957. 
Clausiliidae. 

bermudezae, Bostryx (Pseudoperonaeus) — 
Weyrauch, 1958: 111-112, pl. 9, figs. 38—40; 


WOLFGANG KARL WEYRAUCH 269 


central Peru [HT SMF 156350; PT SMF 
156351, 156352, FML 3114, FMNH 84712]. 
Orthalicidae. 

bicolor, Naesiotus (Naesiotus) — Weyrauch, 
1967a: 408-409, pl. 6, fig. 83; southern Peru 
[HT FML 3987]. Orthalicidae. 

Bilamelliferus — Weyrauch, 1958: 118-119. 
Type-species by original designation: 
Bulimus tschudii Troschel, 1852. Orthalici- 
dae. 

birabenorum, Bostryx (Lissoacme) - 
Weyrauch, 1965a: 71-72, figs. 1-3; 
Tucuman, Argentina [HT FML 985a; PT FML 
985, SMF 164112, FMNH 216893]. 
Orthalicidae. 

cajamarcana, Steeriana (Steeriana) — 
Weyrauch & Zilch, in Zilch, 1954: 73-76, pl. 
5, fig. 8, text-fig. 6; Peru [HT SMF 135516; 
PT SMF 69818, 139782, MNJR HSL-1925, 
FMNH 52338, 52403, SBMNH 125339, 
137905, ZMB 97383, 101649, WW 788]. 
Clausiliidae. 

camachoi, Neopetraeus — Weyrauch, 1967a: 
418—420, pl. 5, figs. 68-70; northern Peru 
[HT FML 1240a; PT FML 1240b, 1245, 1541, 
3985, ANSP 204517, 353175, FMNH, 
SBMNH 83361, SMF 164130, USNM, ZMB 
101777, WW 3985, 3986]. Orthalicidae. 

celendinensis, Drymaeus — Weyrauch, 1956c: 
151-152, pl. 11, figs. 10, 11; northern Peru 
[HT SMF 155307; PT SMF 155308, 69498, 
ANSP 355514, FML 12492, FMNH 53997, 
216831, MCZ 211950, WW 492]. 
Orthalicidae. 

celendinensis, Steeriana (Steeriana) — 
Weyrauch & Zilch, in Zilch, 1954: 70-72, pl. 
5, fig. 6; text-fig. 3; Peru [HT SMF 135517; 
PT SMF 69820, FMNH 52337, 52401, MNRJ 
HSL-1263, SBMNH 137907, ZMB 97382, 
101032, 101648, WW 1366]. Clausiliidae. 

cerrateae, Epiphragmophora (Karlschmidtia) 
— Weyrauch, 1960a: 47-48, pl. 6, fig. 40; cen- 
tral Peru [HT SMF 162030; PT WW 1991]. 
Epiphragmophoridae. 

cerrateae, Hemicena — Weyrauch, 1958: 96-— 
100, pl. 6, fig. 9; text-figs. 1, 2; central Peru 
[HT SMF 140714; PT SMF 139779, 156372, 
ANSP 204510, FEL, FML 1899, 1889b, 
MHN 440, MCZ 202207, 233525, FMNH 
84719, 217050, 217051, SBMNH 125331, 
Universidad Nacional Mayor de San Marcos 
(Lima, Peru), USNM, ZMB 101650, unspeci- 
fied “private collections in U.S.A. and Eu- 
rope”]. Clausiliidae. 

cerrateae, Naesiotus (Raphiellus) [sic = 
Rhaphiellus] — Weyrauch, 1967a: 410-412, 
pl. 5, fig. 77; central Peru [HT SMF 162024; 


PT CAS 80855, FML 1071, 1468]. Ortha- 
licidae. 

chamayensis, Naesiotus (Naesiotus) 
subcostatus — Weyrauch, 1967a: 409-410, 
pl. 7, figs. 97-99; northern Peru. [HT SMF 
162190; PT SMF 162191, CAS 85682, FML 
1078, 3342, FMNH 107830]. Orthalicidae. 

chiletensis, Scutalus (Scutalus) — Weyrauch, 
1967a: 373-375, pl. 2, figs. 24-30; northern 
Peru. [HT FML 1236a (originally WW 1354a); 
PT FML 1236, ANSP 355509, FMNH 54002, 
193423, MNRJ 4416, SBMNH 83313, 
125348, SMF 69811, 181656, 208437, 
277329, USNM, ZMB 97391]. Orthalicidae. 

chusgonensis, Bostryx (Bostryx) — Weyrauch, 
1960a: 30-31, pl. 3, figs. 16, 17; northern 
Peru [HT SMF 162013; PT SMF 162014, 
FML 1855, FMNH 56712]. Orthalicidae. 

cleliae, Zilchogyra — Weyrauch, 1965b: 124, 
pl. 7, fig. “2” [sic = fig. 3 (Zilch, 1970: 236)]; 
Buenos Aires, Argentina [HT FML 977; PT 
FML 10654, MRCN 1091, 1645, SMF 
164142]. Punctidae; = Paralaoma servils 
(Shuttleworth, 1852) (Hausdorf, 2002: 127- 
128). 

combinai, Mesembrinus (Ornatimormus) — 
Weyrauch, 1958: 135-136, pl. 8, fig. 16; cen- 
tral Peru [HT SMF 156201; PT WW 1535]. 
Oleacinidae. 

compactus, Bostryx (Bostryx) zilchi — 
Weyrauch, 1960c: 125, pl. 12, figs. 27, 28; 
central Peru [HT SMF 162156; PT SMF 
162157, FML 3356, FMNH 216887]. 
Orthalicidae. 

costatus, Bostryx (Bostryx) pygmaeus — 
Weyrauch, 1960c: 122, pl. 11, figs. 12-16; 
central Peru [HT SMF 162098; PT SMF 
162099, 162100, ANSP, BMNH, FML 3318, 
FMNH 107850, MCZ 233541, МММ, USNM]. 
Orthalicidae. 

costifer, Bostryx (Elatibostryx) imeldae — 
Weyrauch, 1960c: 129, pl. 12, figs. 35-38; 
central Peru [HT SMF 162101; PT SMF 
162102, 162103, AMS, ANSP, BMNH, FML 
3319 (ex IML 1107), MCZ 233535, FMNH 
107828, 216903, 216904, NMC, NNM, 
SBMNH 83310, USNM,]. Orthalicidae. 

costulatus, Scutalus (Vermiculatus) — 
Weyrauch, 1967a: 395-396, pl. 3, fig. 45; cen- 
tral Peru [HT SMF 162068; PT SMF 162069, 
FML 1230, 1454, 3338, FMNH 107850, 
SBMNH 83316, USNM]. Orthalicidae. 

crassicostata, Temesa (Temesa) decimvolvis 
— Weyrauch, 1958: 102, pl. 6, fig. 2; central 
Peru [HT SMF 156231; PT FML 3122, FEL, 
ЕММН 217006, MCZ 233550, USNM]. 
Clausiliidae. 


270 BARBOSA ET AL. 


crenulatus, Lopesianus — Weyrauch, 1958: 
121, pl. 6, figs. 7, 8; Brazil [HT SMF 156356 
(incorrectly as 156376 by Weyrauch (1958) 
and by Zilch (1970)); PT SMF 155708, FML 
1898, FMNH 216993]. Orthalicidae. 

crucilineatus, Bostryx (Peronaeus) - 
Weyrauch, 1967a: 361-362, pl. 1, fig. 14; 
central Peru [HT FML 1241]. Orthalicidae. 

cuencaensis, Temesa (Neniatracta) adusta — 
Weyrauch, 1964c: 152, figs. 7, 8; central 
Peru [HT SMF 156217; PT SMF 156218, 
FML 1041]. Clausiliidae. 

cuzcoensis, Scutalus (Vermiculatus) — 
Weyrauch, 1967a: 396-398, pl. 4, figs. 47, 
48; southern Peru [HT FML 1225 (ex WW 
10522a) (may not be separated from 
paratype); PT FML 1225 (ex WW 10522), 
SMF 164124, Angel E. Florez collection 
(Cuzco, Peru)]. Orthalicidae. 

cylindricus, Bostryx (Pseudoperonaeus) — 
Weyrauch, 1960c: 127-128, pl. 11, fig. 3; 
central Peru [HT SMF 162107; PT FML 
3333, ЕММН 11331, USNM]. Orthalicidae. 

debilisculptus, Scutalus (Scutalus) coraeformis 
— Weyrauch, 1967a: 376-377, pl. 2, figs. 20— 
23; pl. 9, 137, 138; northern Peru [HT FML 
1984 (ex WW 1964a); PT FML 1984 (ex WW 
1964b-g), FMNH 216786, SMF, USNM]. 
Orthalicidae. 

debilisculptus, Thaumastus (Thaumastiella) 
occidentalis — Weyrauch, 1960a: 30, pl. 3, 
fig. 15; northern Peru [HT SMF 162029; PT 
SMF 162082, FML 1630, ANSP, FMNH 
107841, 216807, 216880, MCZ 233545, 
МММ, USNM]. Orthalicidae. 

decimvolvis, Temesa (Temesa) — Weyrauch, 
1957: 21-22, pl. 1, fig. 13; central Peru [HT 
SMF 156215; PT MHN 159, FMNH 217004, 
WW 1532]. Clausiliidae. 

dedicata, Andinia (Ehrmanniella) — Weyrauch 
& Zilch, in Zilch, 1954: 68-70, pl. 5, fig. 5, 
text-fig. 5; Peru [HT SMF 135515; PT SMF 
69816, 139781, FMNH 52334, 52404, 
217041, 217045, SBMNH 83357, WW 
1414]. Clausiliidae. 

densestrigatus, Mesembrinus (Ornatimormus) 
henrypilsbryi — Weyrauch, 1958: 134-135, 
pl. 8, fig. 20; central Peru [HT SMF 156293]. 
Orthalicidae. 

Diaphanomormus [subgenus of Drymaeus] — 
Weyrauch, 1964b: 57. Type species by origi- 
nal designation: Drymaeus (Diaphano- 
mormus) coelestini obesus Weyrauch, 
1964b, non Martens, 1893, = D. (Mesem- 
brinus) pseudobesus Breure, 1979: 123. 
Orthalicidae. 


Elatibostryx [subgenus of Bostryx] - 
Weyrauch, 1958: 112. Type-species by origi- 
nal designation: Bostryx (Elatibostryx) 
imeldae Weyrauch, 1958. Orthalicidae. 

elegantulus, Naesiotus — Weyrauch, 1956b: 
45, pl. 1, fig. 1; northern Peru [HT SMF 
162023; PT SMF 157276, ANSP 194996, 
CAS 64901, FML 1305, FMNH 53993, MNH 
55, SBMNH 35316, ZMB 97385, 101457]. 
Orthalicidae. 

eliseoduartei, Systrophia (Scolodonta) — 
Weyrauch, 1966: 46-47, fig. 6; Uruguay [HT 
FML 10686a; PT FML 10686, Museo de 
Historia Natural Montevideo, Uruguay]. 
Scolodontidae. 

fernandezae, Naesiotus (Maranhoniellus) — 
Weyrauch, 1958: 122-123, pl. 9, figs. 45, 46; 
northern Peru [HT SMF 157277; PT SMF 
157278, 157530, ANSP 290048, FML 3077, 
CAS 64233, FMNH 84718, SBMNH 83311, 
125519, USNM, unspecified “private collec- 
tions in U.S.A.”]. Orthalicidae. 

flavilabrum, Drymaeus (Mormus) expansus — 
Weyrauch, 1967b: 484-485, pl. 3, fig. 29; 
central Peru [HT FML 1197]. Orthalicidae. 

florezi, Floreziellus — Weyrauch, 1967b: 489- 
490, pl. 1, figs. 7-17; southeastern Peru [HT 
FML 10671a; PT FML 10671, FMNH 
217042, 217044, SMF 164134, Angel E. 
Florez collection (Cuzco, Peru)]. Replaced 
with Bostryx cunyacensis by Breure (1978: 
68—70), who considered this species to be- 
long in Bostryx and thus a junior homonym 
of B. florezi (Weyrauch, 1967a), originally 
named in Phenacotaxus. Orthalicidae. 

florezi, Happia (Happia) — Weyrauch, 1965a: 
75-76, fig. 6; southern Peru [HT FML 
10644a; PT FML 10644, SMF 164140, An- 
gel E. Florez collection (Cuzco, Peru)]. 
Scolodontidae. 

florezi, Incania — Weyrauch, 1964b: 41-43, fig. 
3; southern Peru [HT FML 1218]. Clausi- 
liidae. 

florezi, Phenacotaxus (Ataxellus) — Weyrauch, 
1967a: 369-371, pl. 4, figs. 49, 50; south- 
ern Peru [HT FML 10522a; PT FML 10522b, 
10745, SMF 164151]. Orthalicidae. 

florezi, Radiodiscus — Weyrauch, 1965c: 105- 
106, fig. 1; southern Peru [HT WW 10468a; 
PT: IML 1215, SMF 164118, USNM, WW 
10468]. Charopidae. 

florezi, Systrophia (Scolodonta) — Weyrauch, 
1967a: 433—435, pl. 6, fig. 86; southern Peru 
[HT FML 10621a; PT FML 10621, SMF 
164133, Angel E. Florez collection (Cuzco, 
Peru)]. Scolodontidae. 


WOLFGANG KARL WEYRAUCH 271 


Floreziellus — Weyrauch, 1967b: 488—489. 
Type species by original designation: 
Floreziellus florezi Weyrauch, 1967b. Con- 
sidered by Breure (1978: 70) to be a syn- 
onym of Bostryx. Orthalicidae. 

franzi, Zilchogyra — Weyrauch, 1965c: 112- 
114, fig. 4; Buenos Aires, Argentina [HT FML 
1217]. Charopidae. 

geophilus, Naesiotus (Naesiotus) — Weyrauch, 
1967b: 477—479, pl. 3, figs. 26-28; northern 
Peru [HT FML 1074a; PT FML 10745, 3343, 
CAS 80866, SMF 164032, WW 4139]. 
Orthalicidae. 

giganteus, Radiodiscus — Weyrauch, 1958: 
105-106, pl. 9a, fig. 51; central Peru [HT 
SMF 156342]. Charopidae. 

globosus, Bostryx (Lissoacme) — Weyrauch, 
1967a: 355-357, pl. 1, figs. 11, 12; south- 
east of Lima, Peru [HT FML 1226 (ex WW 
3972a); PT SMF 164125, FML 1227 (ex WW 
10667), FMNH 216905, WW 3973]. 
Orthalicidae. 

glomeratus, Bostryx (Bostryx) zilchi — 
Weyrauch, 1960c: 124-125, pl. 12, figs. 29— 
34; central Peru [HT SMF 162104; PT SMF 
162105, 162106, 277323, AMS, ANSP, 
BMNH, FML 3320, FMNH 107848, 216888, 
MCZ 233538, MNRJ 4445, NMC, USNM]. 
Orthalicidae. 

golbachi, Radiodiscus — Weyrauch, 1965c: 
106-108, fig. 2; Tucumän, Argentina [HT FML 
724a; PT FML 744, 751, 1281]. Charopidae. 

gracilis, Bostryx (Scansiocohlea) — Weyrauch, 
1967a: 360-361, pl. 1, fig. 1; central Peru 
[HT FML 1102; PT FML 3059, FMNH, SMF 
162133, USNM]. Orthalicidae. 

gracillimus, Naesiotus — Weyrauch, 1956b: 6, 
pl. 1, fig. 3; central Peru [HT SMF 155696; 
PT SMF 155697, ANSP, FML 510, FMNH 
56713]. Orthalicidae. 

grandiportus, Bostryx (Bostryx) bromeliarum 
— Weyrauch, 1958: 109-110, pl. 8, figs. 18, 
19; central Peru [HT SMF 156367; PT SMF 
156374, 156375, FML 5227, FMNH 84709, 
WW 2112]. Orthalicidae. 

grandiportus, Zilchiella — \Weyrauch, 1957: 10- 
13, pl. 1, figs. 5-10; northern Peru [HT SMF 
155710; PT SMF 155711-155713, 156700- 
156702, ANSP 355506, FMNH 57257, 
217049, MCZ 211962, FEL, MNRJ 4460, 
SBMHN 35319, 35320, USNM, ZMB 
101647, WW 2005, unspecified “private col- 
lections in U.S.A. and Europe’). Clausiliidae. 

grandiventris, Scutalus — Weyrauch, 1960a: 
42-43, pl. 5, figs. 27-33; northern Peru [HT 
SMF 155690; PT SMF 155681, 155692, 
155693, 162042, 208440, AMS, BMNH, FML 


1364, FMNH 54003, 125999, MNRJ 4417, 
МММ, NMC, SBMNH 35328, USNM]. 
Orthalicidae. 

granulatus, Scutalus (Scutalus) chiletensis — 
Weyraueh, 1967а: 375, pl. 2, figs: 31,32; 
northern Peru [HT FML 1362a [mixed in with 
paratypes?]; PT FML 1362, FMNH 216787, 
SMF 164127, USNM]. Orthalicidae. 

granulosa, Epiphragmophora — Weyrauch, 
1960а: 46-47, pl. 6, fig. 37; southern Peru 
[HT SMF 162041; PT FML 1632, 2531]. 
Epiphragmophoridae. 

haasi, Bostryx (Bostryx) — Weyrauch, 1960a: 
33-35, pl. 5, fig. 35; near Lima, Peru [HT 
SMF 156370; PT SMF 162083, 162902, 
208028, AMS, ANSP, BMNH, FML 1225, 
1225a-d, FMNH 113322, 193624, MCZ 
211970, NMC, RLM, SBMNH 35329, 
USNM]. Orthalicidae. 

haasi, Columbinia (Pfeifferiella) — Weyrauch, 
1957: 46, pl. 1, figs. 1, 2; northern Peru [HT 
SMF 155715; PT SMF 155716, 155717, 
ANSP 204508, FMNH 57254, 217058, MCZ, 
FEL, WW 2006, SBMNH 35331, 83315, 
USNM]. Clausiliidae. 

haasi, Llaucanianus — Weyrauch, 1967a: 421- 
422, pl. 3, figs. 34-36; northern Peru [HT 
SMF 162046; PT SMF 162047-162049, 
ANSP, FML 1082, FMNH 113308, MCZ, 
NNM, USNM, WW 2008]. Orthalicidae. 

haasi, Naesiotus — Weyrauch, 1956b: 79, pl. 
1, fig. 5, 5a; northern Peru [HT FML 2862b; 
PT FML 2862, ANSP 194995, 195001, CAS 
64903, FMNH 56709, 56710, SMF 153373]. 
Orthalicidae. 

haasi, Systrophia (Systrophia) — Weyrauch, 
1960a: 28, pl. 3, fig. 7; northern Peru [HT 
SMF 162020; PT FML 3067, ЕММН]. 
Scolodontidae. 

henrypilsbryi, Mesembrinus (Ornatimorus) — 
Weyrauch, 1958: 134 [nom. nov. pro 
Drymaeus pilsbryi Weyrauch, 1956c, non 
Zetek, 1934]. Orthalicidae. 

hernandezae, Littoridina — Weyrauch, 1963a: 
251, pl. [1], figs. 1, 2; southern Peru [HT FML 
1001a; PT FML 3908, SBMNH 83358, SMF 
164084, USNM]. Cochliopidae. 

hyltonscottae, Zilchogyra —\Weyrauch, 1965c: 
114—115, fig. 5; Tucuman, Argentina [HT FML 
976]. Charopidae. 

imeldae, Bostryx (Elatibostryx) — Weyrauch, 
1958: 113, pl. 9, fig. 37; central Peru [HT SMF 
156347; PT FML 3115, FMNH 84716]. 
Orthalicidae. 

inflatiportus, Bostryx (Bostryx) obliquiportus — 
Weyrauch, 1960c: 123-124, pl. 12, figs. 22- 
26; central Peru [HT SMF 162144: PT SMF 


272 BARBOSA ET AL. 


162145, 162146, 162184, 162185, 164002, 
ANSP, BMNH, FML 3344, FMNH 107838, 
216882, 216883, 216884, MCZ 233543, 
МММ, USNM]. Orthalicidae. 

isidroensis, Steeriana (Steeriana) 
celendinensis — Weyrauch & Zilch, in Zilch, 
1954: 72-73, pl. 5, fig. 7, text-fig. 4; Peru 
[HT SMF 135518; PT SMF 69821, FMNH 
52336, 52402, ZMB 97384, 101031, WW 
1355]. Clausiliidae. 

klappenbachi, Helicina (Trichohelicina) — 
Weyrauch, 1966: 42-44, figs. 1-3; Misiones, 
Argentina [HT FML 11215a; PT FML 11215]. 
Helicinidae. 

lachayensis, Scutalus (Scutalus) versicolor — 
Weyrauch, 1967a: 383-384, pl. 8, figs. 109- 
115; central Peru [HT FML 147a (evidently 
mixed in with the following paratypes); PT 
FML 147, FMNH 216790, 216791, SMF 
208454, ZMB 101782, WW 4043]. 
Orthalicidae. 

laraosensis, Bostryx (Bostryx) obliquiportus — 
Weyrauch, 1960c: 124, pl. 11, figs. 20, 21; 
central Peru [HT SMF 162187; PT SMF 
162188, 162189, FMNH 107840, 216885, 
FML 3332, USNM]. Orthalicidae. 

laraosensis, Temesa (Temesa) pilsbryi — 
Weyrauch, 1960c: 119-120, pl. 11, fig. 7; 
central Peru [HT SMF 162117; PT SMF 
162118, FEL, FML 3346, MCZ 233527]. 
Clausiliidae. 

latecolumellaris, Naesiotus (Naesiotellus) — 
Weyrauch, 1967a: 415-416, pl. 5, figs. 71, 
72; central Peru [HT FML 1076; PT FML 
3335, FMNH, SMF 162139]. Orthalicidae. 

latestriata, Temesa (Temesa) — Weyrauch, 
1958: 100-102, pl. 6, fig. 1; central Peru [HT 
SMF 156232; PT SMF 156233, FML 1047 
(ex WW 3121), FMNH 84717, 217010, MCZ 
233528, USNM]. Clausiliidae. 

lateumbilicatus, Radiodiscus — Weyrauch, 
1966: 44-45, fig. 4; Mendoza, Argentina [HT 
FML 11005a; PT FML 11005, FMNH 
216225, SMF 164148]. Charopidae. 

Leptomormus [subgenus of Mesembrinus] — 
Weyrauch, 1958: 136-137. Type-species by 
original designation: Drymaeus bequaerti 
Weyrauch, 1956c. Orthalicidae. 

Lilloiconcha — Weyrauch, 1965b: 127. Type- 
species by original designation: Austrodiscus 
superbus tucumanus Hylton-Scott, 1963. 
Charopidae. 

lizarasoae, Bostryx (Pseudoperonaeus) — 
Weyrauch, 1967a: 363-364, pl. 3, figs. 38, 
39; central Peru [HT SMF 162009; PT SMF 
162010, FML 1109, 3130, FMNH, MCZ 
233846, USNM]. Orthalicidae. 


Llaucanianus — Weyrauch, 1967a: 420. Type 
species by original designation: 
Llaucanianus haasi Weyrauch, 1967a. 
Orthalicidae. 

longispira, Bostryx (Pseudoperonaeus) — 
Weyrauch, 1960c: 128, pl. 11, figs. 4, 5; cen- 
tral Peru [HT SMF 162112, PT SMF 162113, 
FML 3354, USNM]. Orthalicidae. 

Lopesianus — Weyrauch, 1958: 120. Type-spe- 
cies by original designation: Lopesianus 
crenulatus Weyrauch, 1958. Orthalicidae. 

macedoi, Scutalus (Vermiculatus) — Weyrauch, 
1967a: 398—400, pl. 3, figs. 42-44; central 
Peru [HT SMF 162070; PT SMF 162071, 
FML 1192, 1452 (ex 2874), 2874, FMNH 
113307, SBMNH 83308, USNM, WW 4053]. 
Orthalicidae. 

mantaroensis, Temesa (Temesa) decimvolvis 
Weyrauch, 1963b: 277-279, pl. 1, fig. 7; cen- 
tral Peru [НТ SMF 156237; PT SMF 156238, 
FML 1045, 3123, FMNH 217007, USNM]. 
Clausiliidae. 

Maranhoniellus [subgenus of Naesiotus] — 
Weyrauch, 1958: 112. Type-species by origi- 
nal designation: Naesiotus pilsbry Weyrauch 
1956b. Orthalicidae. 

marasensis, Scutalus (Vermiculatus) 
cuzcoensis — Weyrauch, 1967a: 398, pl. 2, 
fig. 33; southern Peru [HT FML 10641a; PT 
FML 10641b, SMF 164139, Angel E. Florez 
collection (Cuzco, Peru)]. Orthalicidae. 

maximus, Thaumastus (Quechua) salteri — 
Weyrauch, 1967a: 347-348, pl. 9, fig. 135; 
northern Peru [HT SMF 156381; PT FML 
3202]. Orthalicidae. 

Microbeliscus [subgenus of Obeliscus] — 
Weyrauch, 1964b: 40. Type species by origi- 
nal designation: Obeliscus (Microbeliscus) 
silvaevagus Weyrauch, 1964b. Non Micro- 
beliscus Sandberger, 1875. See Nanno- 
beliscus. Subulinidae. 

minor, Bostryx (Bostryx) haasi — Weyrauch, 
1960a: 35, pl. 5, fig. 34; central Peru [HT 
SMF 156371; PT SMF 162044, ANSP, 
BMNH, FML 1105, 2035 (these two numbers 
may represent a single lot), FMNH 107822, 
MCZ 233547, MNRJ 4452, NNM, NMC, 
SBMNH 80266, USNM]. Orthalicidae. 

minor, Steeriana (Steeriana) celendinensis — 
Weyrauch, 1958: 92, pl. 6, fig. 6; northern 
Peru [HT SMF 156212; PT FML 2095]. 
Clausiliidae. 

minor, Temesa (Temesa) decimvolvis — 
Weyrauch, 1963b: 276-277, pl. 1, figs. 9, 10; 
central Peru [HT FML 1046a; PT FML 
1046b, FMNH 217008, SMF 164036, USNM, 
WW 3274]. Clausiliidae. 


WOLFGANG KARL WEYRAUCH 273 


mirabilis, Epiphragmophora — Weyrauch, 
1960c: 129-130, pl. 12, fig. 42; northern Peru 
[HT SMF 162064]. Epiphragmophoridae. 

mirandoi, Potamopyrgus — Weyrauch, 1963a: 
244-247, pl. [1], figs. 3, 4; northern Peru 
[Hydrobiidae. HT FML 1000a; PT FML 
10455, SBMNH 83360, SMF 164083, 
USNM]. Rissooidea: Hydrobiidae? 

Multifasciatus [subgenus of Bostryx] — 
Weyrauch, 1958: 116-117. Type-species by 
original designation: Bulimus subroseus L. 
Pfeiffer, 1869. Orthalicidae. 

multiguttatus, Drymaeus (Ornatimorus) — 
Weyrauch, 1964b: 55-57, fig. 8; central Peru 
[HT FML 1200 (ex WW 10236)]. Ortha- 
licidae. 

Naesiotellus [subgenus of Naesiotus] — 
Weyrauch, 1967a: 414—415. Type species by 
original designation: Naesiotus (Naesiotellus) 
“columellaris”; sic = latecolumellaris 
Weyrauch, 1967a. Orthalicidae. 

Nannobeliscus [subgenus of Obeliscus] — 
Weyrauch, 1967b: 458. Мот. nov. pro 
Microbeliscus Weyrauch, 1964, non 
Microbeliscus Sandberger, 1875. Subu- 
linidae. 

obesus, Drymaeus (Diaphanomormus) 
coelestini — Weyrauch, 1964b: 58, fig. 11; 
central Peru [HT FML 1196a; PT FML 1196b 
(ex WW 10234), SMF 164025, ? 162119]. 
Non Drymaeus sulfureus obesus (Martens, 
1893) = D. pseudobesus Breure, 1978: 123. 
Orthalicidae. 

obesus, Neopetraeus arboriferus Weyrauch, 
1967a: 416-417, pl. 5, figs. 62, 63; Peru [HT 
FML 1244a; PT FML 1244b, 1448]. 
Orthalicidae. 

obliquiportus, Bostryx (Bostryx) — Weyrauch, 
1958: 110, pl. 9, figs. 35, 36; central Peru 
[HT SMF 156353; PT SMF 156354, FML 
3118, FMNH 84714]. Orthalicidae. 

occidentalis, Thaumastus (Thaumastiella) — 
Weyrauch, 1960a: 28-30, pl. 3, figs. 13, 14; 
northern Peru [HT SMF 162026; PT SMF 
162027, 162028, 208392, ANSP 204515, 
FMNH 53991, 216808, MCZ 211967, USNM, 
ZMB 101463, WW 1346]. Orthalicidae. 

omissus, Scutalus (Vermiculatus) —\Weyrauch, 
1967a: 400—403, pl. 8, figs. 116-120; cen- 
tral Peru [HT SMF 155586; PT SMF 208445, 
FML 1193, 12207 (originally WW 207), 
FMNH 216785]. Orthalicidae. 

omissa, Temesa (Temesa) — Weyrauch, 1957: 
18-21, pl. 1, fig. 4; central Peru [HT SMF 
140715; PT SMF 62569, 62671, 89491, 
156230, ANSP, FMNH 52335, 217011, 
217012, 217013, MCZ 233529, FEL, 


SBMNH 35317, MNRJ 4460, USNM, ZMB 
101645, WW 277, 277A-B]. Clausiliidae. 
orcesi, Mesembrinus (Mormus) expansus — 
Weyrauch, 1958: 130, pl. 7, fig. 15; Ecuador 
[HT SMF 156292; PT FML 3193 (two lots 
with the same number listed from two sta- 

tions)]. Orthalicidae. 

orcesi, Thaumastus (Thaumastus) - 
Weyrauch, 1967b: 473-474, pl. 1, fig. 2; 
northeast of Quito, Ecuador [HT FML 3165; 
PT SMF 156325, Escuela Politénica, Quito, 
Ecuador]. Orthalicidae. 

Orcesiellus [subgenus of Plekocheilus] — 
Weyrauch, 1967b: 468—469. Type species 
by original designation: Plekocheilus 
(Orcesiellus) tenuissimus Weyrauch, 1967b. 
Orthalicidae. 

ormeai, Epiphragmophora — Weyrauch, 
1956c: 158-159, pl. 11, fig. 19; northern Peru 
[HT SMF 155315]. Epiphragmophoridae. 

Ornatimorus [subgenus of Mesembrinus] — 
Weyrauch, 1958: 131-132. Type-species by 
original designation: Drymaeus angulobasis 
Pilsbry, 1944. Orthalicidae. 

ortizi, Bostryx (Bostryx) — Weyrauch, 1967a: 
352-353, pl. 1, fig. 13; northern Peru [HT 
FML 1126]. Orthalicidae. 

ortizpuentei, Scutalus (Scutalus) — Weyrauch, 
1967a: 378-379, pl. 7, fig. 100; northern 
Peru [HT FML 1234 (originally WW 10647)]. 
Orthalicidae. 

palizae, Solaropsis (Psadariella) — Weyrauch, 
1956c: 160-161, pl. 11, figs. 15-17; north- 
ern Peru [HT SMF 155311; PT FML 1536, 
FMNH 57253]. Pleurodontidae. 

Pampasinus [subgenus of Bostryx] - 
Weyrauch, 1958: 113-114. Type-species by 
Original designation: Bostryx (Platybostryx) 
weyrauchi Pilsbry, 1944. Orthalicidae. 

paucistrigatus, Neopetraeus arboriferus — 
Weyrauch, 1967a: 417-418, pl. 5, figs. 65- 
67; northern Peru [HT FML 1239a; PT FML 
1239, 1540, FMNH, SMF 164129, WW 
1540]. Orthalicidae. 

peiranoi, Littoridina — Weyrauch, 1963a: 252- 
254, pl. [1], figs. 5-8; Tucuman, Argentina 
[НТ FML 41a; PT FML 41b-d, SBMNH 
83317]. Cochliopidae. 

peruvianus, Pupoides (Pupoides) albilabris — 
Weyrauch, 1960c: 117-119, pl. 11, figs. 1, 
2; central Peru [HT SMF 162160; PT SMF 
162161, 162162, 156378, ANSP, BMNH, 
FMNH 107843, NNM, USNM, WW 3250]. 
Pupillidae. 

peterseni, Zilchogyra — Weyrauch, 1965b: 
125-126, pl. 7, fig. “3” [sic = fig. 2 (Zilch, 
1970: 236)]; northwestern Peru [HT SMF 


274 BARBOSA ET AL. 


165245a; PT SMF 165245, FML 1221]. 
Charopidae. 

Pfeifferiella [subgenus of Columbinia] — 
Weyrauch, 1957: 34. Type-species by origi- 
nal designation: Columbinia (Pfeifferiella) 
haasi Weyrauch, 1957. Clausiliidae. 

pichitacalugaensis, Mesembrinus (Ornati- 
mormus) henrypilsbryi — Weyrauch, 1958: 
135, pl. 8, fig. 17; central Peru [HT SMF 
156338 (incorrectly as 156388 by Weyrauch 
(1958) and Zilch (1970)); PT WW 3110]. 
Orthalicidae. 

pilosus, Scutalus (Vermiculatus) — Weyrauch, 
1967a: 403-404, pl. 3, figs. 40, 41; central 
Peru [HT SMF 162065; PT SMF 162066, 
FML 1413, 1194, ЕММН]. Orthalicidae. 

pilsbryi, Drymaeus — Weyrauch, 1956c: 153- 
154, pl. 11, fig. 7; Peru [НТ SMF 155303; PT 
FML 192a]. Non Zetek, 1834; see 
henrypilsbryi, Mesembrinus (Ornatimorus). 
Orthalicidae. 

pilsbryi, Naesiotus — Weyrauch, 1956b: 67, pl. 
1, fig. 4; northern Peru [HT SMF 155698; 
PT SMF 155699, ANSP 194999, CAS 
64902, FML 7463, FMNH 56711]. 
Orthalicidae. 

pilsbryi, Systrophia (Systrophia) obvoluta — 
Weyrauch, 1958: 106-107, pl. 6, figs. 4, 5; 
central Peru [HT SMF 156344; PT SMF 
156345, 156346, FML 642, ZMB 109019]. 
Scolodontidae. 

pilsbryi, Temesa —Weyrauch, 1956c: 146-148, 
pl. 11, figs. 1-4; central Peru [HT SMF 
155296; PT SMF 155297, 155293, 155299, 
FML 1043 (ex WW 3058), ANSP 204505, 
(?)355505, MCZ 211974, FMNH 57255, 
FEL, SBMNH 35318, 35335, 361319, 
USNM, ZMB 101646]. Clausiliidae. 

planispira, Systrophia (Systrophia) — 
Weyrauch, 1967a: 429-431, pl. 6, fig. 82; 
northern Peru [HT FML 1266a; PT FML 
1266]. Scolodontidae. 

primigenia, Temesa (Temesa) pilsbryi — 
Weyrauch, 1960c: 120-121, pl. 11, fig. 8; 
central Peru [HT SMF 162147]. Clausiliidae. 

Psadariella [subgenus of Solaropsis] — 
Weyrauch, 1956c: 159. Type-species by 
Original designation: Solaropsis (Psadariella) 
palizae Weyrauch, 1956c. Pleurodontidae. 

Pseudoglandina — Weyrauch, 1967b: 485- 
486. Type species by original designation: 
Pseudoglandina agitata Weyrauch, 1967b. 
Amphibulimidae. 

Pseudoperonaeus [subgenus of Bostryx] — 
Weyrauch, 1958: 111. Type-species by origi- 
nal designation: Bostryx (Pseudoperonaeus) 
bermudezae Weyrauch, 1958. Orthalicidae. 


pygmaeus, Bostryx (Bostryx) — Weyrauch, 
1960c: 121-122, pl. 11, figs. 10, 11; central 
Peru [HT SMF 162127; PT SMF 162128, 
FML 3317, FMNH 107845]. Orthalicidae. 

pygmaea, Temesa (Temesa) albocostata — 
Weyrauch, 1963b: 272-273, pl. 1, figs. 3, 4; 
central Peru [HT SMF 162111; PT SMF, FML 
1039, 3321, FMNH 107829, 217002, USNM, 
WW 3321]. Clausiliidae. 

quadritaeniatus, Drymaeus (Orodrymaeus) 
farrisi — Weyrauch, 1956c: 150-151, pl. 11, 
fig. 9; northern Peru [HT SMF 155306; PT 
WW 2132]. Orthalicidae. 

rehderi, Bostryx (Elatibostryx) — Weyrauch, 
1960a: 35-36, pl. 3, figs. 4, 5; central Peru 
[HT SMF 156386; PT SMF 156387, 156388, 
AMS, ANSP, BMNH, FML 3207, FMNH 
84715, MCZ 233540, MUSNM, WW 3207, 
NNM, NRJ 4447, SBMNH 83309]. 
Orthalicidae. 

rodriguezae, Bostryx (Bostryx) — Weyrauch, 
1967b: 475-476, pl. 3, figs. 33-48; central 
Peru [HT FML 1125a; PT FML 3315, FMNH 
107835, 216918, MNRJ 4439, SBMNH 
83318, SMF 162123, 277316, WW 3331]. 
Orthalicidae. 

scotophilus, Bostryx (Bostryx) — Weyrauch, 
1967a: 353-354, pl. 1:85: 10; central Peru 
[HT FML 1130; PT FML 1424 (presumably 
ex WW 1130)]. Orthalicidae. 

semiclausa, Epiphragmophora diluta — 
Weyrauch, 1960a: 45, pl. 6, fig. 36; south- 
ern Peru [HT SMF 162040; PT FML 1627]. 
Non Hylton-Scott, 1951. See next entry. 
Epiphragmophoridae. 

semiaperta, Epiphragmophora diluta — 
Weyrauch, 1964a: 169 (nov. nom. pro E. 
diluta semiclausa Weyrauch, 1960a, non 
Hylton-Scott, 1951). Epiphragmophoridae. 

shutcoénsis, Тетеза (Temesa) pilsbryi — 
Weyrauch, 1960c: 119, pl. 11, fig. 6; central 
Peru [HT SMF 162114; PT SMF 162115, 
FML 1051, FMNH 107837, WW 3347, 
USNM]. Clausiliidae. 

silvaevagus, Naesiotus — Weyrauch, 1960a: 
36-37, pl. 4, fig. 20; central Peru [HT SMF 
162001; PT FML 3279]. Orthalicidae. 

silvaevagus, Obeliscus (Microbeliscus) — 
Weyrauch, 1964b: 40-41, fig. 2; central Peru 
[HT FML 1219]. Subulinidae. 

similis, Littoridina — Weyrauch, 1963a: 256, pl. 
[1], figs. 13, 14; southern Peru [Hydrobiidae. 
HT FML 1002a; PT FML 1002b, 3883, 
SBMNH 83359, SMF 164090, USNM]. 
Cochliopidae. 

souzalopesi, Drymaeus (Drymaeus) - 
Weyrauch, 1965a: 73-74, figs. 4, 5; Goyaz, 


WOLFGANG KARL WEYRAUCH 215 


Brazil [HT WW 10622a; PT WW 10622, 
FMNH 216828, 216990]. Orthalicidae. 

subterranea, Columbinia (Pfeifferiella) — 
Weyrauch, 1957: 68, pl. 1, fig. 3; northern 
Peru [HT SMF 155718; PT SMF 156234, 
FMNH 217059, MCZ 233530, WW 2007]. 
Clausiliidae. 

superba, Peruinia flachi — Weyrauch, 1960a: 
24-25, pl. 3, figs. 1-3; central Peru [HT SMF 
156235; PT SMF 156236, 162036, 162037, 
FEL, FLM 3092, FMNH 107866, 217018, 
МММ, USNM, Rolf A. Brandt collection (now 
in the SMF)]. Clausiliidae. 

superbus, Bostryx (Multifasciatus) — 
Weyrauch, 1967a: 365-367, pl. 1, figs. 15- 
19; central Peru [HT FML 1137a (ex WW 
3330); PT FML 1137, FMNH 107866, 
217018, SMF 162120]. Orthalicidae. 

tarmensis, Naesiotus (Reclasta) — Weyrauch, 
1967b: 479-480, pl. 4, figs. 51, 52; central 
Peru [HT SMF 162017, PT SMF 162018, 
FML 4146, 4147, CAS 80848, FMNH 
216991, MNRJ 4418, WW 1459]. Ortha- 
licidae. 

tenuissimus, Plekocheilus (Orcesiellus) — 
Weyrauch, 1967b: 469-470, pl. 2, fig. 23, 
pl. 4, fig. 50; Pichincha, Ecuador [HT WW 
3364]. Orthalicidae. 

terrestris, Steeriana (Cylindronenia) maran- 
honensis — Weyrauch, 1964b: 44—45, figs. 
4, 5; northern Peru [HT FML 702a; PT FML 
702, ANSP, FMNH 30733, SMF 62566, 
62677]. Clausiliidae. 

Thaumastiella [subgenus of Thaumastus] — 
Weyrauch, 1956b: 11-12. Type-species by 
original designation: Bulimulus (Protoglyptus) 
sarcochrous Pilsbry, 1897. Orthalicidae. 

thomei, Bulimulus (Rhinus) — Weyrauch, 
1967b: 481-482, pl. 1, figs. 3-5; Rio Grande 
do Sul, Brazil [НТ МКСМ 1021a; PT MRCN 
1021, FML 1262, FMNH 217442, WW 
10669]. Orthalicidae. 

thomei, Radiodiscus — Weyrauch, 1965b: 121- 
122, pl. 7, fig. 1; southeastern Brazil [HT 
MRCN 1073a; PT MRCN 1073, FML 1262]. 
Charopidae. 

translucidus, Drymaeus (Drymaeus) — 
Weyrauch, 1967a: 422-424, pl. 6, figs. 78, 
79; central Peru [HT FML 1202a; PT FML 
1202b, SMF 162158]. Orthalicidae. 

Trichohelicina [subgenus of Helicina] — 
Weyrauch, 1966: 41-42. Type species by 
original designation: Helicina ( Trichohelicina) 
klappenbachi Weyrauch, 1966. Helicinidae. 

tridentata, Zilchistrophia — Weyrauch, 1960а: 
27-28, pl. 3, fig. 6; central Peru [HT SMF 
162006; PT FML 3280]. Family uncertain. 


Trochogyra [subgenus of Zilchogyra] — 
Weyrauch, 1965b: 126. Type-species by 
original designation: Endodonta superba 
Thiele, 1927. Charopidae. 

turritus, Naesiotus (Raphiellus) [sic = 
Rhaphiellus] — Weyrauch, 1967a: 412-414, 
pl. 5, fig. 64, 64a; northern Peru [HT FML 
1229a (ex WW 3976a); PT FML 1229 (ex 
WW 3976)]. Orthalicidae. 

Unilamellatus [subgenus of Ptychodon] — 
Weyrauch, 1965b: 134. Type-species by 
original designation: Ptychodon (Uni- 
lamellatus) unilamellatus Weyrauch, 1965b. 
Endodontidae. 

unilamellatus, Ptychodon (Unilamellatus) — 
Weyrauch, 1965b: 134, pl. 7, fig. 4; central 
Peru [HT WW 3863a; PT SMF 164138, WW 
3863]. Endodontidae. 

Vermiculatus [subgenus of Scutalus] - 
Weyrauch, 1967a: 384—405. Unavailable; no 
differentiation description or definition pro- 
vided and no type species designated (ICZN 
Code Art. 13); first made available by Breure 
(1978: 166); type species: Bulinus bicolor G. 
B. Sowerby |, 1835. Orthalicidae. 

vilchezi, Bostryx (Bostryx) —Weyrauch, 1960a: 
32-33, pl. 3, figs. 8-10; northern Peru [HT 
SMF 155704; PT SMF 155705, 162038, 
162039, 208048, 277322, AMS, ANSP 
204513, BMNH, FML 2009, FMNH 84711, 
216897, 216899, MCZ 211969, NNM, NMC, 
MNRJ 4441, SBMNH 35330, 361320, 
USNM]. Orthalicidae. 

willinki, Bostryx (Bostryx) — Weyrauch, 1964b: 
54-55, fig. 12; Catamarca, Argentina [HT 
FML 121a; PT FML 121]. Orthalicidae. 

woytkowskii, Leptarionta — Weyrauch, 1960c: 
130-131, pl. 11, fig. 9; northern Peru [HT 
SMF 164000; PT WW 3358]. Xanthony- 
chidae. 

wygodzinskyi, Radiodiscus — Weyrauch, 
1965с: 110-112, fig. 3; Tucuman, Argentina 
[HT FML 4148a; PT: FML 764, 1282, WW 
4148]. Charopidae. 

zilchi, Bostryx (Bostryx) — Weyrauch, 1958: 
108—109, pl. 9, figs. 41, 42; central Peru [HT 
SMF 156348; PT SMF 156349, FML 1185 
(ex WW 3113), FMNH 216886, MCZ 233539, 
USNM]. Orthalicidae. 

zilchi, Epiphragmophora — Weyrauch, 1960a: 
44—45, pl. 6, figs. 41-44; southern Peru [HT 
SMF 156391; PT SMF 156392, 156393, 
156394, FML 1428, FMNH 56707, W. Biese 
collection (Santiago, Chile)]. Epiphragmo- 
phoridae. 

zilchi, Naesiotus — Weyrauch, 1956b: 910, pl. 
1, fig. 6; northern Peru [HT WW 1908; PT 


276 BARBOSA ET AL. 


ANSP 194998, ЕММН 56714, SMF 155701]. 
Orthalicidae. 

zilchi, Scutalus (Vermiculatus) culmineus — 
Weyrauch, 1967a: 393-394, pl. 9, figs. 139, 
140; central Peru [HT FML 3328a; PT FML 
3328, 4032, SMF 162141]. Orthalicidae. 

zilchi, Stenostylus — Weyrauch, 1956c: 156- 
157, pl. 11, fig. 18; Peru [HT SMF 155314]. 
Orthalicidae. 

zilchi, Systrophia (Systrophia) — Weyrauch, 
1967a: 431-433, pl. 6, fig. 81; southern Peru 
[HT SMF 162007; PT SMF 162008, FML 
1273, 2108, 4001, ЕММН]. Scolodontidae. 

zilchi, Temesa (Temesa) — Weyrauch, 1963b: 
285-288, pl. 1, fig. 14; central Peru [HT FML 
1053; PT FML 3329, SMF 162177, USNM]. 
Clausiliidae. 

Zilchiella — Weyrauch, 1957: 9. Type-species 
by original designation: Zilchiella grandi- 
portus Weyrauch, 1957. Clausiliidae. 

Zilchistrophia — Weyrauch, 1960a: 26. Type- 
species by original designation: Zilchistrophia 
tridentata Weyrauch, 1960a. Family uncertain. 

Zilchogyra — Weyrauch, 1965b: 122-123. 
Type-species by original designation: Helix 
costellata d’Orbigny, 1836. Charopidae. 

zischkai, Nenia (Columbinia) — Weyrauch, 
1956a: 113-115, pl. 6, fig. 3, 3a; eastern 
Bolivia [HT WW 1368; PT ANSP 328089]. 
Clausiliidae. 


MOLLUSCAN TAXA NAMED FOR 
WEYRAUCH 


Bostryx weyrauchi Pilsbry, 1944. The Nauti- 
lus, 57(3): 87-88, pl. 9, fig. 5 [Gastropoda]. 

Neopetraeus weyrauchi Pilsbry, 1944. The 
Nautilus, 57(3): 88, pl. 9, fig. 4 [Gastropoda.]. 

Thaumastus (Scholvienia) weyrauch [sic] 
Pilsbry, 1944. The Nautilus, 57(4): 121, pl. 
11, fig. 2, 2a [Gastropoda]. 

Nenia weyrauchi Pilsbry, 1945. The Nautilus, 
58(3): 82, pl. 3, fig. 5 [Gastropoda]. 

“Austroselenites (?)” weyrauchi Haas, 1951. 
Fieldiana, Zoology, 31(46): 535-537, fig. 119 
[Gastropoda]. 


ACKNOWLEDGEMENTS 


We thank the following curators for informa- 
tion on the type collections in their institutions: 
Maria Gabriela Cuezzo, Instituto Miguel Lillo, 
Argentina; Rina Ramirez, Museo de Historia 
Natural, Universidad Nacional Mayor de San 


Marcos, Peru; Arnaldo C. dos Santos Coelho, 
Museu Nacional, Universidade Federal do Rio 
de Janeiro, Brasil; Jochen Gerber, Field Mu- 
seum of Natural History, Chicago, Illinois, 
U.S.A.; Robert Hershler, United States National 
Museum, Smithsonian Institution, Washington, 
D.C., U.S.A.; Ronald Janssen, Naturmuseum 
Senckenberg, Germany; Frank Kohler, Mu- 
seum für Naturkunde, Humboldt-Universitat, 
Berlin, Germany; Christina Piotrowski, Califor- 
nia Academy of Sciences, San Francisco, Cali- 
fornia; U.S.A.; Gary Rosenberg, Academy of 
Natural Sciences of Philadelphia, Pennsylva- 
nia, U.S.A.; Luiz Ricardo L. de Simone, Museu 
de Zoologia, Universidade de Sao Paulo, Brasil; 
and Paul Valentich-Scott, Santa Barbara Mu- 
seum of Natural History, Santa Barbara, Cali- 
fornia, U.S.A. Gerardo Lamas, Museo de 
Historia Natural, Universidad Nacional Mayor 
de San Marcos, Peru; Virginia Sanchez Puerta, 
Indiana University, Bloomington, Indiana, 
U.S.A.; José Ochoa, Peru; Jose Willibaldo 
Thomé, Pontificia Universidade Catolica do Rio 
Grande do Sul, Brasil; and Eduardo 
Vasconcelos, Museu Nacional, Universidade 
Federal do Rio de Janeiro, Brasil, helped with 
biographical and bibliographical information. 
Rudiger Bieler, Field Museum of Natural His- 
tory, Chicago, Illinois, U.S.A.; Ligya dos Reis 
Correa, Instituto Oswaldo Cruz, Brazil; Kaspar 
Delhey, Max Planck Institut for Ornithology, 
Radolfzell, Germany; Richard E. Petit, North 
Myrtle Beach, South Carolina, U.S.A.; and Barry 
Roth, San Francisco, California, U.S.A., pro- 
vided advice or reviewed the manuscript. Alan 
R. Kabat, Washington, D.C., U.S.A., looked up 
type information in the Museum of Compara- 
tive Zoology, Harvard University, Cambridge, 
Massachusetts, U.S.A. 


LITERATURE CITED 


AGUILAR, P., 1970, Prof. Dr. Wolfgang K. 
Weyrauch, 1907-1970. Revista Peruana de 
Entomologia, 13: 3—4. 

BOUCHET, P. & J.-P. ROCROI, 2005, Classifi- 
cation and nomenclator of gastropod families. 
Malacologia, 47(1-2): 397 pp. [classification of 
Caenogastropoda by W. Ponder & P. Bouchet 
and of Pulmonata by B. Hausdorf & P. Bouchet]. 

BREURE, А. $. H., 1978, Notes on and descrip- 
tion of Bulimulidae (Mollusca, Gastropoda). 
Zoologische Verhandelingen Uitgegeven door 
het Rijksmuseum van Natuurlijke Historie te 
Leiden, 164: 255 pp., 22 pls. 

BREURE, A. S. H., 1979, Systematics, phylog- 
eny and zoogeography of Bulimulinae (Mol- 


WOLFGANG KARL WEYRAUCH Zee 


lusca). Zoologische Verhandelingen Uitgegeven 
door het Rijksmuseum van Natuurlijke Historie 
fe Leiden, 168: 215 pp., 3 pls. 

DRAHG, F. & M. G. CUEZZO, 1999, Catalogo de 
especimenes tipo de la colecciön malacolögica 
de la Fundacion Miguel Lillo. Acta Zoolögica 
Lilloana, 45(1): 55-65. 

DUARTE, E., 1970, Noticia del fallecimiento del 
gran malacologo Dr. Wolfgang K. Weyrauch. 
Comunicaciones de la Sociedad Malacolögica 
del Uruguay, 3(19): 13-14. 

HAUSDORF, B., 2002, Introduced land snails 
and slugs in Colombia. Journal of Molluscan 
Studies, 68(2): 127-131. 

INTERNATIONAL COMMISSION ON ZOOLOGI- 
CAL NOMENCLATURE, 1999, International 
Code of Zoological Nomenclature, 4" ed. Lon- 
don, -T.Z.N. xxix + 306 pp. 

KOHLER, F., 2007, Annotated type catalogue of 
the Bulimulinae (Pulmonata, Orthalicoidea, 
Bulimulidae) in the Museum für Naturkunde 
Berlin. Mitteilungen aus dem Museum für 
Naturkunde in Berlin, Zoologische Reihe 83(2): 
125-159. 

LAMAS, G., 1981, Historia de la entomologia en 
el Peru Il: periodo de los viajeros, colectores y 
estudiosos especializados. Revista Peruana de 
Entomologia, 23(1): 25-31. 

MEYER, T. & W. K. WEYRAUCH, 1965a, Guia 
para excursiones biolögicas а los arededores 
de la ciudad de Tucuman. Departamento de 
Extension Universitaria Nacional de Tucuman, 
44 pp. 

MEYER, T. & W. K. WEYRAUCH, 1965b, Guia 
para dos excursiones biolögicas en la Provincia 
de Tucumán, 2™ ed., Miscelania 23, Instituto 
Miguel Lillo. 127 pp. 

NEUBERT, E. & R. JANSSEN, 2004, Die Typen 
und Typoide des Natur-Museums Senckenberg, 
84: Mollusca: Gastropoda: Pulmonata: 
Orthalicoidea: Bulimulidae (2), Orthalicidae, 
Placostylidae. Archiv für Molluskenkunde, 
133(1/2): 193-297, incl. 24 pls. 

WEYRAUCH, W. K., 1956a [May 10], Two new 
species of Clausiliidae from Peru and Bolivia. 
The Nautilus, 69(4): 110-115, pl. 6. 

WEYRAUCH, W. K., 1956b [June 20], The ge- 
nus Naesiotus, with descriptions of new spe- 
cies and notes on other Peruvian Bulimulidae. 
Proceedings of the Academy of Natural Sci- 
ences of Philadelphia, 108: 1-17, pl. 1. 

WEYRAUCH, W. K., 1956c [November 1], Neue 
Landschnecken aus Peru. Archiv für 
Molluskenkunde, 85(4/6): 145-162, pl. 11. 

WEYRAUCH, W. K., 1957 [May 31], Sieben neue 
Clausiliiden aus Peru. Archiv für Mollusken- 
kunde, 86(1/3): 1-28, pl. 1. 

WEYRAUCH, W. K., 1958 [December 1], Neue 
Landschnecken und neue Synonyme aus 
Südamerika, 1. Archiv für Molluskenkunde, 
87(4/6): 91-140. 

WEYRAUCH, W. K., 1960a [June 30], Zwanzig 
neue Landschnecken aus Peru. Archiv ftir 
Molluskenkunde, 89(1/3): 23-48, pls. 3-6. 

WEYRAUCH, W. K., 1960b [June 30], Zur 
Kenntnis von Newboldius (Bulimulidae). Archiv 
für Molluskenkunde, 89(1/3): 49-56, pls. 7-8. 


WEYRAUCH, W. K., 1960c [December 5], Sieb- 
zehn neue Landschnecken aus Peru. Archiv für 
Molluskenkunde, 89(4/6): 117-132, pls. 11-2. 

WEYRAUCH, W. K., 1963a [June 10], Cuatro 
nuevas especies de Hydrobiidae de Argentina 
y Peru (Gastropoda, Prosobranchia). Acta 
Zoologica Lilloana, 19: 243-259, pl. 1. 

WEYRAUCH, W. K., 1963b [June 10], Aporte al 
conocimiento de Temesa, | (Clausiliidae, Mol- 
lusca). Acta Zoologica Lilloana, 19: 261-288, 
Bl. 1. 

WEYRAUCH, W. K., 1964a [July 20], Nomenkla- 
torische Bemerkungen. Archiv für Mollusken- 
kunde, 93(3/4): 169. 

WEYRAUCH, W. K., 1964b [December 30], 
Nuevos gastropodos terrestres y nuevos 
sinönimos de Sudamerica. Il. Acta Zoologica 
Lilloana, 20: 33-60. 

WEYRAUCH, W. K., 1964c [December 30], 
Aporte al conocimiento de Temesa, ll 
(Clausiliidae, Mollusca). Acta Zoologica 
Lilloana, 20: 145-162, pl. 1. 

WEYRAUCH, W. K., 1965a [August 1], Tres 
nuevos gastrópodos terrestres de Sudamérica. 
Neotropica, 11(35): 71-76. 

WEYRAUCH, W. K., 1965b [September 30], 
Neue und verkannte Endodontiden aus Süd- 
amerika. Archiv fur Molluskenkunde, 94(3/4): 
121-134, pl. 7. 

WEYRAUCH, W. K., 1965c [December 1], Cinco 
nuevos Endodontidos de Argentina y Peru 
(Gastropoda, Euthyneura). Neotropica, 11(36): 
105-115. 

WEYRAUCH, W. K., 1966 [August 1], Gastro- 
podos terrestres de Argentina, Uruguay y 
Brasil. Neotropica, 12(38): 41-47. 

WEYRAUCH, W. K., 1967a [November], Treinta 
y ocho nuevos gastropodos terrestres de Peru. 
Acta Zoologica Lilloana, 21: 343-455, pls. 1-9. 

WEYRAUCH, W. K., 1967b [November], 
Descripciones y notas sobre gastropodos 
terrestres de Venezuela, Colombia, Ecuador, 
Brasil y Peru. Acta Zoologica Lilloana, 21: 457- 
499, pls. 1-4. 

WEYRAUCH, W. K., 1967c [December 22], 
Microhappia Thiele (1927) ist ein synonym von 
Punctodiscops H. B. Baker (1925) (Gas- 
tropoda, Systrophiidae). Archiv fur Mollusken- 
kunde, 96(3/6): 139-141. 

WEYRAUCH, W. К. & Р. CORONADO, 1958, 
Lugares en las cercanias de Lima, mas 
apropiados para las excursiones de Ciéncias 
Biologicas, con alumnus de educacion 
secundaria. |.- Las Lomas. Imprenta Colegio 
Nac. Ntra. Sra. de Guadalupe, 17 pp. 

WILLINK, A., 1999, Biografias Lilloanas. Revista 
de la Sociedad Entomolögica Argentina, 58(3- 
4): 3-10. 

ZILCH, A., 1954 [March 15], Landschnecken aus 
Peru, 2. Archiv fur Molluskenkunde, 83(1/3): 
65-79, pls. 5-6. 

ZILCH, A., 1970, Wolfgang Karl Weyrauch (1907- 
1970). Mitteilungen der Deutschen Malako- 
zoologischen Gesellschaft, 2(18), 226-236. 


Revised ms. accepted 10 January 2008 


‘Gu ЭР Mer: 
ee 


58 his 4 ‚Аа МА ee = 
их Ro 
ART ИБО DOTES ls 
LE пресной: +5 ЗЕ: Seta 


seed 


si alse D 
“hi pr sx 


ты ue Ge is 
| знати о re": a 
+ RRR es te Saude ial cab E 

41 GE и uh ; ees 
BAT À. au FOGARTY a 
у EC Bg wie + rl a 


| + ind erat: KR ap | 

mee erde» ein ei 
ig se Bert Je b Zhe 
de q eN We A, a 


tis 


went ro À m a yr rares sn. 


oti neared tobera ha Math: 
ae asada OE: e | we ae 

ere BER HO SEDE À, ALS 
SMe оком es bal DATE 
BIETE 


are 


nahi a 
te ee anyone Ayo A St NAS $ мат 
pad mal deci da at 


AT ne 


7 ae Stee ЗЕЕ 
nist A ah ave 
= sects | 


RNA PA HIS E 


CRT CT TR er tu TT 
Shea” a ae a ee ey A EE Zr 
a Gas Dé i u ТЕ ow Ar Le Mis us A 
‘| A = 2 on! "pa | ‚м fi thy | 7 + к A Opes pu as et re si ur 

Fr =F Eee, tpg US = a | a i Et nl NA pa ЗИ. 

Ds ER pes AZ ! Sh Ae I m. Van ee Е 

en Er м Е ea D | EE, 
A Md E г.“ al Ah, 

Ра pi 4 ar = a 
ie A ety 
À EN nm 
HR LEN; 
Deer 


$ Aare PBs 


$ 
мы tie 
la dl sl 
As Вы Г. 
=i) ig 


Es cr = LAR 4 
3 Er = es МА 
ES aye! ; Ag 


O 
oe ae LE | 


ke E 


‘a ¿e 


ane, ES Alen e er caer cri 
tg Li y A Ze 
rag ЕС “he a E ‘i a и, i A * 
В e we F р Te Ace Li : 
м and ss PT ee 2% er Tee 


: > io ACA ВН 
A ra а 
Ber ee re a 

pi: v 4 œ ANS RD 7 


ui: eR BEN: és 


i D 


ыы ee, Г 


he ech À: ne he 
LES fie een | 


свеч: ай caf 5 и. = 
ENR Panda и ser 
nen nae Бы 


(ESE e | “on =. 

FE a Kg | a Mab aa 
= Pee роса Fr TE vers de : Е des Y a 
po a FA fn УР HH. = roe > SEX SS La 


pa | ä Le po va pr 
pe Sethe: sy Pape none % RR 
Le Woks A. 4) A FE TA ars ARGENT Kae 
OS ar À a ААУ Wy 
‘aie wen: 4 АЙ va 


rh 21 mit. 
Pr Zsa SS 


„авик а metio ey pone tE ag 


ie a‘ Be Bala Sines | M + ~ zu Ves а ve 


ER a MO 4, RENE 
oe 


oy A ER ER un 


+f A 

# а cet). 

pl VE EU ie AA FEES DER 

POMC er ая 
ni SEE OB E ES A oa 


MALACOLOGIA, 2008, 50(1-2): 279-292 


ULTRASTRUCTURAL STUDIES OF OOGENESIS AND SEXUAL MATURATION 
IN FEMALE CHLAMYS (AZUMAPECTEN) FARRERI FARRERI 
(JONES & PRESTON, 1904) (PTERIOMORPHIA: PECTINIDAE) 
ON THE WESTERN COAST OF KOREA 


Ee-Yung Chung 


School of Marine Life Science, Kunsan National University, 
Gunsan 573-701, Korea; eychung@kunsan.ac.kr 


ABSTRACT 


Ultastructural studies of the development and degeneration of the oocytes and follicle 
cells in female Chlamys (Azumapecten) farreri farreri (Jones & Preston, 1904) are de- 
scribed for scallops collected from Daehuksando, Jeollanam-do, Korea. Vitellogenesis 
occurred by way of endogenous autosynthesis and exogenous heterosynthesis. Auto- 
synthesis involved the combined activity of the Golgi complex, mitochondria, and rough 
endoplasmic reticulum, whereas heterosynthesis involved endocytotic incorporation of 
extraovarian precursors at the basal region of the early vitellogenic oocytes prior to the 
formation of the vitelline coat. Auxiliary cells were involved in the development of the 
previtellogenic and early vitellogenic oocytes and appear to play an integral role in vitello- 
genesis and oocyte degeneration by assimilating products originating from the degener- 
ated oocytes, thus allowed the transfer of yolk precursors needed for vitellogenesis. Auxiliary 
cells presumably have a lysosomal system for breakdown products of oocyte degenera- 
tion. The reproductive cycle in females was classified into five stages: Stage |: early active 
stage (January to March), Stage Il: late active stage (March to April), Stage Ill: ripe stage 
(April to August), Stage IV: partially soawned stage (June to August), and Stage V: spent/ 
inactive stage (August to January). The spawning period continued from June to August, 
with a peak between July and August when the seawater temperature was exceeded 22°C. 
The percentage of first sexual maturity was 59.3% in individuals of 50.1-60.0 mm in shell 
height, and 100% in those > 70.1 mm in shell height. 

Because harvesting clams less than 50.1 mm in shell height could potentially cause a 
drastic reduction in recruitment, a measure indicating a prohibitory fishing size should be 
enacted for adequate fisheries management. 

Key words: Chlamys (Azumapecten) farreri farreri, oogenesis, auxiliary cell, first sexual 


matration. 


INTRODUCTION 


The Jicon scallop, Chlamys (Azumapecten) 
farreri farreri (Jones 8 Preston, 1904) is a com- 
mercially important bivalve in East Asian coun- 
tries, including Korea, Japan, and China. On 
the western coast of Korea, this species is 
mainly found in the gravel bed in the subtidal 
zone at depths up to 10 m (Yoo, 1976; Kwon 
et al., 1993; Min et al., 2004). Due to past over- 
harvesting, it has been identified as a species 
requiring a more sustainable fishing regimen. 
For the propagation and management of this 
species, it is important that we fully understand 
the reproductive biology with regard to germ 
cell differentiation during oogenesis and sexual 


219 


maturation. Previously there have been many 
studies on reproduction in C. farreri farreri, in- 
cluding aspects of the reproductive cycle (Lioa 
et al., 1983; Yakovlev & Afeichuk, 1995), growth 
and spawning (Na et al., 1995; Kang € Zhang, 
2000), experimental triploids and tetraploids 
(Yang et al., 1999a), its distribution and ecol- 
ogy (Whang & Kim, 1973), larval growth 
(Kuang et al., 1997; Yang et al., 1999b), and 
experimental aquaculture (Lim et al., 1995; Sun 
et al., 1996, 1997). 

Despite this, there are still significant gaps 
in our knowledge regarding its reproductive 
biology. Above all, studies on the development 
and degeneration of the oocytes and auxiliary 
cells during oogenesis of C. farrerii farreri are 


280 CHUNG 


34° 38' М 


34° 36' М 


125° 24'Е 


125° 30'Е 


ЕС. 1. Мар of the sampling area. 


required to understand this animal’s reproduc- 
tive biology. In the majority of bivalve species, 
the ovaries contain auxiliary cells, a kind of 
accessory cell, that play a role in the storage, 
mobilization, and synthesis of yolk precursors 
during oogenesis (Wourms, 1987). More spe- 
cifically, oocyte degeneration, which is known 
as atresia, is a commonly observed phenom- 
enon in most bivalve species. In bivalves, the 
products of lysis material created by the auxil- 
lary cells act as sources of metabolites that can 
be rapidly mobilized by the organism (Pipe, 
1987; Dorange et al., 1989; Le Pennec et al., 
1991; Gaulejac et al., 1995). Above all, the func- 
tions of the auxiliary cells in the resorption of 
the lysis products of atretic oocytes of this spe- 
cies should be investigated in further detail. 
Understanding of the reproductive cycle and 
spawning period of this species will provide 
information needed for the determination of the 
size at first reproduction and the recruitment 
period. Additional information on the shell size 
attained when 50% of the individuals reach 
first sexual maturity can determine a prohibi- 
tory size for adequate natural resource man- 


agement. Therefore, the purpose of this paper 
is to describe vitellogenesis during oogenesis, 
the reproductive cycle, and the size at first 
sexual maturity in C. farreri farreri, using cyto- 
logical, histological, and morphometric proce- 
dures. Results will be useful for improved 
fisheries management of this species. 


MATERIALS AND METHODS 
Sampling 


Jicon scallops were collected monthly by 
dredge in the gravel bed in the subtidal zone 
at depths up to 10 m off Daehuksando Island, 
Jeollanam-do, Korea (Fig. 1), from January 
2003 to December 2004. 


Histology (Light Microscopy) 


For light microscopic examination of histo- 
logical preparations, female ovarian tissues 
were removed from animals and preserved in 
Bouin’s fixative for 24 h, then washed with run- 


OOGENESIS & SEXUAL MATURATION IN CHLAMYS FARRERI 281 


ning tap water for 24 h. Tissues were then de- 
hydrated in alcohol and embedded in paraffin 
molds. Embedded tissues were sectioned at 
5-7 um thickness using a rotary microtome. 
Sections were mounted on glass slides, 
stained with Hansen hematoxylin — 0.5% 
eosin, and examined using a light microscope. 
These stained sections were analyzed to (1) 
describe the ovarian cycle, and (2) determine 
the size at which sexual maturity is attained, 
the methods described below. 


Ovarian Cycle by Light Microscopical Obser- 
vation 


To describe the ovarian cycle and to identify 
the spawning period, a total of 731 ovarian 
histological preparations were made from scal- 
lops of 50.1-94.7 mm shell height. 


Ultrastructure of Germ Cells and Auxiliary Cells 
During Oogenesis and Oocyte Degeneration 


A total of 95 females was used for ultrastruc- 
tural study of germ cells and auxiliary cells by 
electron microscopy. For transmission electron 
microscopy, excised samples of gonads were 
cut into small pieces and fixed immediately in 
2.5% paraformaldehyde-glutaraldehyde in 
0.1 M phosphate buffer (pH 7.4) for 2 h at 4°C. 
After prefixation, the specimens were washed 
several times in the buffer solution and then 
postfixed in 1% osmium tetroxide solution in 
0.2 M phosphate buffer (pH 7.4) for 1 h at 4°C. 
Specimens then were dehydrated in increasing 
concentrations of ethanol, cleared in propylene 
oxide, and embedded in an Epon- Araldite mix- 
ture (Epon-812). Ultrathin sections of Epon- 
embedded specimens were cut to a thickness 
of 80-100 nm with a LKB ultramicrotome. The 
sections were mounted on collodion-coated 
copper grids, double stained with uranyl acetate, 
followed by lead citrate, and observed under a 
JEM 100 CX-2 (80 kv) electron microscope. 


Size at First Sexual Maturity by Light Micro- 
scopical Observation 


For determination of the size at 50% of first 
sexual maturity, a total of 208 ovarian histo- 
logical preparations (30.4-94.7 mm shell 
height) were examined the size at 50% first 
sexual maturity (= biological minimum size) by 
histological observations from May to Octo- 
ber 2003. The percentage (%) of first sexual 
maturity = No. of mature individuals x 100/No. 
of total individuals investigated. 


RESULTS 
Position and Morphology of the Ovary 


Chlamys farreri farreri is dioecious. The ovary 
is conical or crescent shaped, and it is sepa- 
rated from the digestive diverticula and the 
adductor muscle. It is located from the ventral 
region of the visceral mass to the adductor 
muscle. The ovary is a diffuse organ composed 
of highly branching follicles (acini) in which 
germ cells develop. 

As ovarian maturation progressed, the ovary 
encircled part of the adductor muscle, and the 
external color of the ovary became pink (the 
testis being milky white or light yellow). There- 
fore, sex could be easily determined from ex- 
ternal features. At this time, mature oocytes 
readily emerged when the ovary was slightly 
scratched. After spawning, the ovary degen- 
erated, and then the sexes became difficult to 
distinguish. 


Annual Reproductive Cycle with Ovarian De- 
velopmental Stages 


Based on electron microscopical and histo- 
logical observations of the germ cells and other 
surrounding cells (auxiliary cells), the gonadal 
phases were classified into five successive 
stages (Fig. 2). The stages and the criteria 
used in defining them are as follows: 


Stage 7 (early active stage): Oogonia and 
previtellogenic oocytes propagate along the 
follicular wall of the ovary. The oogonia were 
about 10-11 um in diameter, and the pevi- 
tellogenic oocytes 16-20 um in diameter. 
The lumina of the oogenic follicles were 
empty during the early active stage, although 
the auxiliary cells, which were attached to 
the previtellogenic oocyte, appeared in the 
oogenic follicle at this stage (Fig. 3A). In 2003 
and 2004, female individuals in stage | (early 
active stage) appeared from January to 
March when seawater temperatures were 
about 10°C. 


Stage Il (late active stage): At a size of 30- 
40 um in diameter, each early vitellogenic 
oocyte formed an egg-stalk connected to the 
follicular wall (germinal epithelium), and the 
auxiliary cells, which were attached to the 
oocyte, appeared in the lumen of the follicle. 

At a diameter of 40-50 um, each late 
vitellogenic oocyte had a large germinal 
vesicle and an egg-stalk attached to the fol- 


NO 
00 
N 


W. TEMP. (*C) 


30 

20 

10 

0 
100 == |= 
| == 
90 ee el 
Sa Е 
| Е ЕЕ 
80 SEN ie) === 
| Coo ES] == 
< 70 DEEE 
| = 
о 60 Е 
2 LEZ 
E 50 D mil 
ка Е 
> 40 CO HA E 
fabs CE 
2 | HE 
E 30 Se a 
| A= 
20} "o 
10 | ВЕНЕ 
BEBE Е 

2 S 


Ш Early Active stage 


CHUNG 


==) (IZ 


SM 


J FMAMJJ A 


MONTH 


Late Active stage 


M Ripe stage 


Partially spawned stage Е Spent/Inactive stage 


FIG. 2. Frequency of gonadal phases in female Chlamys farreri farrericompared with mean seawater 


temperatures from January 2003 to December 2004. 


licular wall (Fig. 3B). In 2003 and 2004, fe- 
male individuals in stage II (late active stage) 
were found from March to April when seawa- 
ter temperatures were relatively low (7-13°C). 


Stage Ill (ripe stage): The majority of matur- 
ing oocytes grew to 50-60 um in diameter, 
becoming round or oval in shape, and were 
located in the center of the lumen. Each ripe 
ovum (60-70 um in diameter) was sur- 
rounded by a gelatinous membrane and its 
cytoplasm was filled with a large number of 
yolk granules (Fig. 3C). At this time, the aux- 
iliary cells detached from the mature oocyte. 
In 2003 and 2004, female individuals in stage 
Ill (ripe stage) appeared from April through 
August when seawater temperature gradu- 
ally increased over 15°C. 


Stage IV (partially soawned stage): Most ripe 
ova were discharged from the oogenic fol- 
licles, although a few undischarged mature 
oocytes as well as vitellogenic oocytes re- 
mained (Fig. 3D). In 2003 and 2004, female 
individuals in stage IV (partially spawned 
stage) were found from June to August, and 
the main spawning occurred between July 
and August when seawater temperatures 
were higher than 22°C. 


Stage V (spent/inactive stage): After spawn- 
ing, each follicle contracted and degener- 
ated, and the undischarged oocytes in the 
lumen of the follicle underwent cytolysis. The 
products of gamete atresia were resorbed 
(Fig. 3E). Thereafter, a rearrangement of the 
connective tissues was observed. Occasion- 


OOGENESIS & SEXUAL MATURATION IN CHLAMYS FARRERI 283 


FIG. 3. Photomicrographs of oogenic follicles in various gonadal phases in female Chlamys farreri 
farreri. A: Early active stage; В: Late active stage; С: Ripe stage; D: Partially spawned stage; E: Spent 
stage; F: Inactive stage. Abbreviations: AC = auxiliary cell; DO = degenerated oocyte; ES = egg stalk; 
EVO = early vitellogenic oocyte; FW = follicular wall; LU = lumen; LVO = late vitellogenic oocyte; MO = 
maturing oocyte; OG = oogonium; PVO = previtellogenic oocyte; RO = ripe ovum; UDO = undis- 
charged oocyte. 


284 CHUNG 


FIG. 4. Electron micrographs of oogenesis in female Chlamys farreri farreri. A: Oogonia, with a large 
nucleus, several mitochondria, and vacuoles in the cytoplasm; B: Previtellogenic oocyte, with a nucleo- 
lus in the nucleus, the mitochondria, rough endoplasmic reticulum, and an attached auxiliary cell 
containing a nucleus, mitochondria, and rough endoplasmic reticulum; C: Early vitellogenic oocyte 
and desmosome (arrow head), with a nucleus and several lipid droplets, and attached auxiliary cells 
containing lipid droplets and glycogen particles; D: Early vitellogenic oocyte, with the Golgi products 
in the Golgi complex near lipid droplets. Abbreviations: AC = auxiliary cell; DS = desmosome; EVO = 
early vitellogenic oocyte; G = Golgi complex; GP = glycogen particle; GPR = Golgi product; LD = lipid 
droplet; M, mitochondrion; N = nucleus; OG = oogonium; PVO = previtellogenic oocyte; RER = rough 
endoplasmic reticulum; SER = smooth endoplasmic reticulum; V = vacuole. 


OOGENESIS & SEXUAL MATURATION IN CHLAMYS FARRERI 285 


ally, a few oogonia were present on the folli- 
cular walls in this stage (Fig. 3F). In 2003 
and 2004, female individuals in stage (spent/ 
inactive stage) appeared from August 
through January. 


Ultrastructure of Germ Cells and Auxiliary Cells 
during Oogenesis and Oocyte Degeneration 


Based on electron microscopical observa- 
tions, four distinct phases of oogenesis were 
distinguished in germ cells, that is, oogonia, 
previtellogenic oocytes, vitellogenic oocytes, 
and mature oocytes (Eckelbarger & Davis, 
1996). 


Oogonia: The oogonia, which measured from 
10-11 um in diameter, were round in shape. 
Each possessed a large ovoid nucleus, in 
which the chromatin was reticular and mar- 
ginal. The oogonia, which multiply within the 
follicular wall (germinal epithelium), were 
single of formed a cluster in the follicle. Sev- 
eral mitochondria, the Golgi complex, and 
vacuoles appeared in the cytoplasm of оо- 
gonia (Fig. 4A). 


Previtellogenic Oocytes: The oogonia develop 
into previtellogenic oocytes. At the begining 
of cytoplasmic growth of the previtellogenic 
oocyte, several mitochondria and vacuoles 
were concentrated around the nucleus. At 
this time, the auxiliary cells initially appeared 
close to the oocyte, and thereafter, progres- 
sively surrounded the oocyte. Close contact 
was maintained with the auxiliary cell. Near 
the adherence zone, vacuoles were visible 
in the cytoplasm of the auxiliary cells (Fig. 
4B). 


Vitellogenic Oocytes: In the early vitellogenic 
oocyte, lipid droplets, mitochondria, and en- 
doplasmic reticulum were usually present in 
the perinuclear region. Desmosomes were 
found in the attached parts of the early 
vitellogenic oocyte connected to the auxil- 
iary cell. In particular, the mitochondria, lipid 
droplets, and glycogen particles appeared 
in the cytoplasm of the auxiliary cells (Fig. 
4C). During early oogenesis, lipid droplets 
appeared near the Golgi product formed by 
the Golgi complex in the cytoplasm of the 
early vitellogenic oocyte (Fig. 4D), and were 
also found between the mitochondria and 
well-developed rough endoplasmic reticulum 
in the cytoplasm of the early vitellogenic oo- 
cyte (Fig. 5A). At this time, coated vesicles, 


resulting from endocytosis appeared at the 
basal region of the early vitellogenic oocyte. 
The uptake of nutritive material in the coated 
vesicles formed by receptor-mediated en- 
docytosis appeared through the formation of 
coated endocytotic pits on the oolemma (Fig. 
9B). At the same time yolk granules clearly 
appeared among the mitochondria, lipid 
droplets, and the endoplasmic reticulum at 
the cortical region (Fig. 5C). In the mid- 
vitellogenic oocyte, multivesicular bodies, 
which were formed by modified mitochon- 
drial cristae, appeared between the nuclear 
envelope and the cortical region (Fig. 5D). 
Thereafter, yolk granules and lipid droplets 
filled the cytoplasm of the late vitellogenic 
oocyte, whereas the auxiliary cells gradually 
lost their intimate association with the late 
vitellogenic oocyte surface. The cytoplasm 
of the auxiliary cells, which was detached 
from the oocyte, was filled with vacuoles and 
myelin figure (Fig. 6A). After yolk granules 
were mixed with coated vesicles (by endocy- 
tosis) and multivesicular bodies, and then 
small yolk granules were formed near the 
cortical region of the late vitellogenic oocyte 
(Fig. 6B). In the late stages of oogenesis, 
large yolk granules were formed by a com- 
bination of small yolk granules (Fig. 6C). 


Mature Oocytes: In the mature oocyte, small 


yolk granules were continuously combined 
and became larger mature yolk granules in 
the cytoplasm. A mature yolk granule in com- 
posed of three components: (1) crystalline 
core, (2) electron lucent cortex, and (3) a lim- 
iting membrane (Fig. 6D). 


Oocyte Degeneration: The degenerating оо- 


cytes appeared sligtly irregular or polyhedral 
near the auxiliary cells and were deformed 
by compression in the follicle. A number of 
vacuoles, degenerating yolk granules a few 
phagosomes (lysosomes), and lipid droplets 
appeared in the cytoplasm of degenerating 
oocyte. At this stage, especially in the auxil- 
lary cells, a few phagosomes (lysosomes) and 
a number of vacuoles, and a small number of 
lipid droplets appeared in the cytoplasm of 
the auxiliary cell, whereas glycogen particles 
decreased in the cytoplasm of the auxiliary 
cells, which were attached to the degenerat- 
ing oocyte (Fig. 7A). During the gradual dis- 
integration of the oocytes, the endoplasmic 
reticulum was specifically involved in the de- 
generative process. The smooth or rough 
endoplasmic reticulum became distended, 


286 CHUNG 


FIG. 5. Electron micrographs of oogenesis in female Chlamys farreri farreri. A: Early vitellogenic 
oocyte, with lipid droplets between well-developed rough endoplasmic reticulum and the mitochon- 
dria; B: Early vitellogenic oocyte, with coated vesicles at the basal region through the coated endocy- 
totic pits (upper left) formed by endocytosis; C: Early vitellogenic oocyte, with yolk precursors among 
rough endoplasmic reticulum, mitochondria, and lipid droplets; D: Mid-vitellogenic oocyte, with 
multivesicular bodies formed by the modified mitochondria. Abbreviations: CP = coated pit; CV = 
coated vesicle; EVO = early vitellogenic oocyte; LD = lipid droplet; M = mitochondrion; MM = modified 
mitochondrion; MVB = multivesicular body; MVO = mid-vitellogenic oocyte; OL = oolemma; RER = 
rough endoplasmic reticulum; YG = yolk granule. 


OOGENESIS & SEXUAL MATURATION IN CHLAMYS FARRERI ZO 


FIG. 6. Electron micrographs of oogenesis in female Chlamys farreri farreri. A: Late vitellogenic oocyte, 
with a number of yolk precursors, lipid droplets, microvilli on the vitellogenic coat, and auxiliary cells 
containing lipid droplets, vacuoles, and a myelin figure; B: Late vitellogenic oocyte, with cortical gran- 
ules near the vitelline coat, and proteinaceous yolk granules; C: Late vitellogenic oocyte, with proteina- 
ceous yolk granules and a number of immature yolk granules; D: Mature oocyte, with a number of 
mature yolk granules each composed of three parts: 1) crystalline core, 2) electron lucent cortex, and 3) 
a limiting membrane. Abbreviations: AC = auxiliary cell; CC = crystalline core; CG = cortical granule; 
ELC = electron lucent cortex; LD = lipid droplet; LM = limiting membrane; LVO = late vitellogenic oo- 
cyte; M = mitotochondrion; MF = myelin figure; MO = mature oocyte; MV = microvillus; MYG = mature 
yolk granule; N = nucleus; PHA = phagosome; V = vacuole; VC = vitelline coat; YG = yolk granule. 


288 


CHUNG 


FIG. 7. Electron micrographs of degenerated oocytes with auxiliary cells in 
female Chlamys farerri. farreri. A: Degenerating oocyte, with degenerating yolk 
granules, phagosomes by lysosome, and the auxilliary cells containing lipid 
droplets and various phagosomes and vacuoles in the cytoplasm; B: Degener- 
ated oocyte, with distended endoplasmic reticulum, vacuoles, degenerated 
granules, myelin figure organelle, phagosomes (lysosomes) near the lipid drop- 
lets, and abnormal vitelline coat. Abbreviations: AC = auxiliary cell; AMS = 
abnormal microvillus structure; DER = distended endoplasmic reticulum; DGR 
= degenerating granule; DO = degenerated oocyte; LD = lipid droplet; LM = 
limiting membrane; LY = lysome; M = mitochondrion; MF = myelin figure; N = 
nucleus; PHA = phagosome; V = vacuole; VC = vitelline coat. 


which led to vacuolation of the ooplasm. At 
this time, abnormal microvillus structure ap- 
peared on the vitelline coat of the degener- 
ated oocyte. Mitochondria and yolk granules 
disintegrated in the ooplasm, and lysis was 
initiated at the cell periphery, several vacu- 


oles and numerous heterogenous, dense 
granules that appear similar to phagosomes 
(lysosomes) were present in the ooplasm. In 
particular, many disintegerated granules with 
myelin figure and phagosomes were visible 
at the periphery of the oocyte (Fig. 7B). 


OOGENESIS & SEXUAL MATURATION IN CHLAMYS FARRERI 


209 


TABLE 1. Shell height at first sexual maturity in female Chlamys (Azumapecten) 
farreri farreri from May to October 2003. Ind. = Individual. 


Shell height Number of Individuals by Gonadal Stage* Total Mature 
(mm) EA LA RI PS SP/IA Ind. (%) 
30.4—40.0 24 24 0 
40.1-50.0 15 6 2 2 28 39.3 
50.1-60.0 11 4 8 4 27 59,3 
60.1-70.0 2 2 17 13 4 38 94.7 
70.1-80.0 16 12 4 32 100.0 
80.1-90.0 1S 9 6 30 100.0 
90.1-94.7 14 10 5 29 100.0 

Total 208 


*Gonadal stage: EA = Early Active Stage; LA = Late Active Stage; RI = Ripe Stage; PS = Partially 


Spawned Stage; SP/IA = Spent/Inactive Stage. 


Size at First Sexual Maturity 


As shown in Table 1, it was found that go- 
nadal development of smaller individuals rang- 
ing from 30.4—40.0 mm in shell height were in 
the early active stage, characterized by a small 
number of oogonia and the previtellogenic 
oocytes present. During the period between 
June and August, when spawning was ob- 
served among older individuals. However, 
younger animals (30.4 to 40.0 mm in shell 
height) had a small number of oogonia and a 
number of previtellogenic oocytes were 
present in the follicles of the ovary. It is sup- 
posed that their sizes at sexual maturity could 
not have been reached until late August when 
spawning was completed. In addition, the per- 
centage of first sexual maturity of female scal- 
lops ranging from 40.1-50.0 mm shell height 
is 39.3%, but those individuals were in a vari- 
ety of gonadal stages during the breeding sea- 
son. In contrast, all individuals of shell height 
greater than 70.1 mm displayed ripe, partially 
spawned, or spent/inactive stages. Accord- 
ingly, it is assumed that most individuals can 
reach full maturity by late August if they are 
larger 70.1 mm in shell height at that time. 


DISCUSSION 
Gamete Differentiation and Vitellogenesis 
Although many authors suggested the for- 
mation of lipid droplets in several species, no 


clear morphological evidence has been shown 
for the processes involved in lipid droplet for- 


mation thus far (Pipe, 1987; Dorange & Le 
Pennec, 1989; Gaulejac et al., 1995). In our 
present study, however, lipid droplets appeared 
among the Golgi complex, well-developed en- 
doplasmic reticulum, and mitochondria in the 
early vitellogenic oocytes. Therefore, it is as- 
sumed that they may be involved in the forma- 
tion of lipid droplets (Chung & Ryou, 2000; 
Chung et al., 2002, 2005, 2006; Chung, 2007). 
Vitellogenesis showed a possibility of auto- 
synthetic and heterosynthetic yolk formation. 

The yolk granules originate in the cortical 
regions of the oocyte, and then fill the entire 
ooplasm of the oocyte. However, the sizes of 
yolk granules varies in different regions of the 
egg. Therefore, various cell organelles, in par- 
ticular the Golgi complex, endoplasmic reticu- 
lum and mitochondria are thought to be 
involved in endogenous formation of yolk gran- 
ules in the cytoplasm (Pipe, 1987; Dorange & 
Le Pennec, 1989; Gaulejac et al., 1995; Chung 
et al., 2005; Chung, 2007), while the uptake 
of nutritive material in the coated vesicle 
formed by receptor-mediated endocytosis ap- 
peared through the formation of coated en- 
docytotic pits on the oolemma (Chung, 2007). 
From these observations, it is assumed that 
vitellogenesis in С. farreri farreri occurs by way 
of endogeous autosynthesis and exogenous 
heterosynthesis. Autosynthesis involves the 
combined activity of the Golgi complex, mito- 
chondria, and rough endoplasmic reticulum; 
heterosynthesis occurs as the incorporation 
of extraovarian precursors into oocytes by 
endocytosis, which involves the basal region 
of the early vitellogenic oocytes prior to the 
formation of the vitelline coat. 


290 CHUNG 


Functions of the Auxiliary Cells 


In stage | (early active stage), the auxiliary 
cells at the periphery of the oogenic follicle (or 
acinus) initially appears close to the previ- 
tellogenic oocyte, and thereafter, progressively 
surrounds a part of the oocyte. At this stage, a 
small number of vacuoles were visible in the 
cytoplasm of the auxiliary cells near the ad- 
herence zone. The attached auxiliary cells also 
showed cytological modifications as their cy- 
toplasmic volume increased in Crassostrea 
virginica (Eckelbarger & Davis, 1996) and 
Mytilus edulis (Pipe, 1987). 

In the present study of C. farreri farreri, sev- 
eral auxiliary cells attached to previtellogenic 
and early vitellogenic oocytes during the early 
stages of oogenesis, but, most auxiliary cells, 
detached from the mid- or late vitellogenic оо- 
cytes; only a few cells appeared near the stalk 
region of the oocyte. At the adherence zone 
between of the auxiliary cells and vitellogenic 
oocytes, lipid droplets and a number of vacu- 
oles, and the myelin figures appeared in the 
cytoplasm of the auxiliary cells, which is indica- 
tive of membrane breakdown (Pipe, 1987). 
Because the auxiliary cells are abundant on 
the oocyte in the early stages of oogenesis and 
gradually detach from the vitellogenic oocyte, 
it is assumed that auxiliary cells function as 
nutritive cells in the early formation and devel- 
opment of the oocytes. In the scallop, Pecten 
maximus, Dorange & Le Pennec (1989) pro- 
posed that the “auxiliary cells” of P maximus 
might play a role in oocyte nutrition and vitelline 
envelope formation partially due to the pres- 
ence of extensive RER cisternae. RER cister- 
nae have been reported in many bivalve follicle 
cells (Gaulejac et al., 1995), while the follicle 
cells of Crassostrea gigas were reported to 
contain smooth endoplasmic reticulum and 
desmosomes (Eckelbarger & Davis, 1996). In 
this study, smooth endoplasmic reticulum and 
desmosomes between oocyte and auxiliary 
cells could be seen. Therefore, our results co- 
incide with the report of Eckelbarger & Davis 
(1996), and nutrients in the auxiliary cells pre- 
sumably transport to the oocyte by the des- 
mosome. 

Pipe (1987) reported that endocytotic figures 
appeared between vitellogenic oocytes and the 
auxiliary cells, indicating a transfer nutrients in 
Mytilus edulis. In the present study, endocy- 
totic-coated vesicles, indicating a transfer nu- 
trients, appeared between the auxiliary cells 
and the vitellogenic oocytes. Therefore, the 
results mentioned above showed a similar phe- 
nomenon to those reported by Pipe (1987). 


Oocyte Degeneration and the Functions of the 
Auxiliary Cells 


In the present study, the characteristics of a 
functional role of lysosomes and a number of 
degenerated yolk granules containing a few 
myelin figures appeared in the ooplasm of the 
degenerated oocytes in C. farreri farreri. At the 
same time, several phagosomes (lysosomes) 
near the lipid droplets appeared in the cyto- 
plasm of the auxiliary cells, that were attached 
to the degenerated oocytes. In particular, mor- 
phologically similar phagosomes (lysosomes), 
which were easily observed in the cytoplasm 
of degenerated oocytes, also appeared in the 
auxiliary cells. Thus, the auxiliary cells appear 
to play an integral role in vitellogenesis and 
oocyte degeneration. During the period of oo- 
cyte degeneration, the auxiliary cells function 
in phagocytosis and intracellular digestion of 
products originating from oocyte degeneration; 
these cells might also have a function associ- 
ated with the induction of oocyte degeneration, 
and it is assumed that they are also active in 
the resorption of phagosomes (lysosomes) from 
the degenerated oocyte, because lipid droplets 
and degenerating phagosomes appeared in the 
auxiliary cells. In this study, the number of lipid 
granules gradually increased in auxiliary cells 
during gametogenesis; this function can per- 
mit a transfer of yolk precursors necessary for 
vitellogenesis and allow for the accumulation 
of reserves in the cytoplasm as glycogen and 
lipids, which can be employed by vitellogenic 
oocytes (Gaulejac et al., 1995). Therefore, it is 
assumed that the auxiliary cells, that are at- 
tached to degenerated oocytes, presumably 
have a lysosomal system for breakdown of in- 
gested material, and they might be involved in 
the induction of oocyte degeneration, and might 
also resorb various phagosomes (lysosomes) 
in the cytoplasm for nutrient storage, such as 
lipid droplets, during oocyte degeneration, as 
seen in Meretrix lusoria (Chung, 2007). 


Fate of the Gametes 


Regarding reproductive energy allocated to 
the production of gametes, some authors 
(Morvan & Ansell, 1988) stated that continu- 
ous production and resorption of gametes can 
be regarded as an adaptation to environmen- 
tal temperature and food availability. If the en- 
ergy allocated to the production of gametes is 
too large, nutritive reserves might not be suffi- 
cient to allow all eggs to reach the critical size 
and maturity for spawning and fertilization. In 
this case, the products of gamete atresia can 


OOGENESIS & SEXUAL MATURATION IN CHLAMYS FARRERI 2% 


be resorbed and the energy reallocated to still- 
developing oocytes or used for other metabolic 
purposes by marine mollusks (Dorange & Le 
Pennec, 1989; Mortavkine & Varaksine, 1989). 

In Mytilus edulis, after spawning, gamete 
resorption is common in the acini of the ovary. 
It is supposed that Mytilus edulis resorbs ga- 
metes in follicles to utilize the high nutritive 
reserves in developing oocytes for other meta- 
bolic activities (Pipe, 1987) as observed in 
other bivalves (Dorange & Le Pennec, 1989; 
Motavkine & Veraksine, 1989). Therefore, itis 
assumed that C. farreri farreri has a similar 
reproductive mechanism to resorb and utilize 
high nutritive substances rather than releas- 
ing non-viable gametes. 


Size at First Sexual Maturity 


The percentage of first sexual maturity of 
individuals of 50.1 to 60.0 mm in shell height, 
in the late active, ripe, and partially spawned 
stages, was over 50%, and was 100% in those 
over 70.1 mm in shell height in the late active, 
ripe, partially spawned, and spent/inactive 
stages. Accordingly, it is likely that most indi- 
viduals will have reached maturity by late Au- 
gust if larger than 70.1 mm in shell height. This 
means that larger individuals can reach matu- 
rity earlier than smaller individuals, indicating 
that harvesting scallops less than 50.1 mm in 
shell height could cause a drastic reduction in 
recruitment. Accordingly, an enforced, mini- 
mum legal fishing size should prove adequate 
for fisheries management of this species. 
Thus, in particular, information on the size at 
50% of sexual maturity is very important, and 
can determine a prohibitory fishing size for 
adequate natural resources management 
through determination of size at 50% first 
sexual maturity. However, further detailed 
studies on age determination of this species 
should be carried out to better understand the 
population dynamics. 


ACKNOWLEDGEMENTS 


The author is grateful to Dr. William Heard 
of the Florida State University for helpful com- 
ments on the manuscript. Thanks are due also 
to Mr. Ye-Kyu Lee of the Electron Microscope 
Laboratory, Korea University, for his assistance 
with transmission electron microscopy. This 
research was supported in part by a fund from 
the Research Projects (2003-2004) of the 
Fisheries Science Research Institute, Kunsan 
National University. 


LITERATURE CITED 


CHUNG, E. Y., 2007, Oogenesis and sexual 
maturation in Meretrix lusoria (Réding, 1798) 
(Bivalvia: Veneridae) in western Korea. Jour- 
nal of Shellfish Research, 26: 1-10. 

CHUNG, E. \., 5. Y. KIM. К. Я. PARK € GM 
PARK, 2002, Sexual maturation, spawning, and 
deposition of the egg capsules of the female 
purple shell, Rapana venosa (Gastropoda: 
Muricidae). Malacologia, 44: 241-257. 

CHUNG, E. Y., S. Y. KIM, G. M. PARK & J. M. 
YOON, 2006, Germ cell differentiation and 
sexual maturation of the female Neptunea 
(Barbitonia) arthritica cumingii (Crosse, 1862) 
(Gastropoda: Buccinidae). Malacologia, 48: 
65-76. 

CHUNG Е. Y, Vd, PARK, Wl) У. LEE“G ОГК. 
RYU, 2005, Germ cell differentiation and sexual 
maturation of the hanging cultured female scal- 
lop Patinopecten yessoensis on the east coast 
of Korea. Journal of Shellfish Research, 24: 
913-921. 

CHUNG, E. Y. & D. K. RYOU, 2000, Gametoge- 
nesis and sexual maturation of the surf clam 
Mactra veneriformis on the west coast of Ko- 
rea. Malacologia, 42: 149-163. 

DORANGE, G. & M. LE PENNEC, 1989, Ultra- 
structural study of oogenesis and oocytic de- 
generation in Pecten maximus from the Bay of 
St. Brieyc. Marine Biology, 103: 339-348. 

DORANGE, G., Y. M. PAULET 8 M. LE PENNEC, 
1989, Etude cytologique de la partie femelle 
de la gonade de Pecten maximus recolte en 
baie de Saint-Brieu. 2. Ovogenese et lyse 
ovocytaire. Haliotis, 19: 299-314. 

ECKELBARGER, K. J. & C. V. DAVIS, 1996, 
Utrastructure of the gonad and gametogenesis 
in the eastern oyster, Crassostrea virginica. 1. 
Ovary and oogenesis. Marine Biology, 127: 79- 
87 


GAULEJAC, De. B., M. HENRY & N. VICENTE, 
1995, An ultrastructural study of gametogenesis 
of the marine bivalve Pinna nobilis (Linnaeus 
1758). 1. Oogenesis. Journal of Molluscan Stud- 
ies, 61: 375-392. 

KANG, Т. С. & С. 1. ZHANG, 2000, A study on the 
growth and spawning of Korean scallop 
(Chlamys farreri) around Wando, Korea. Jour- 
nal of the Korean Fisheries Society, 36: 210- 
221 [in Korean]. 

KUANG, S., H. SUN, F. LI & J. FANG, 1997, Feed- 
ing and growth of scallop Chlamys farreri be- 
fore and after spawning. Marine Fisheries 
Research of China, 17: 80-86. 

KWON, О. K., С. M. PARK & J. S. LEE, 1993, 
Coloured shells of Korea. Academy Publish- 
ing Company. 288 pp. [in Korean]. 

LE РЕММЕС, M. P. С. BENINGER, С. DORANGE 
& Y. M. PAULET, 1991, Trophic sources and 
pathways to the developing gametes of Pecten 
maximus (Bivalvia: Pectinidae). Journal of the 
Marine Biological Association of the United 
Kingdom, 71: 451-463. 

LIM, H. K., С. $. GO € Y. H. LEE, 1995, Studies 
on the technology development for seed produc- 
tion of Chlamys farreri. Pp. 355-360, in: Techni- 
cal Report of South Sea Regional Fisheries 


292 CHUNG 


Research Institute, National Fisheries Research 
and Development Institute, Yosu, Korea. 

LIAO, C., У. XU & Y. WANG, 1983, Reproductive 
cycle of the scallop Chlamys farreri (Jones and 
Preston) at Qingdao. Journal of Fisheries of 
China, 7: 1-13. 

MIND. Ко LEE. DB. KO'S Jy Gi JEFZ004, 
Mollusks in Korea. Hanguel Graphics, Busan, 
Korea. 566 pp. [in Korean]. 

MORVAN, С. 8 A. D. ANSELL, 1988, Streological 
methods applied to the reproductive cycle of 
Tapes rhomboides. Marine Biology, 97: 355- 
364. 

MOTAVKINE, P. A. & A. A. VARAKSINE, 1989, 
La reproduction chez les mollusques bivalves: 
rôle du systême nerveux et regulation. Reports 
Scientifiques et Techniques de l'IFREMER, 10: 
1-250. 

NA, G. H., W. G. JEONG & C. H. CHO, 1995, A 
study on seedling production of Jicon scallop, 
Chlamys farreri. 1. Spawning, development and 
rearing of larvae. Journal of Aquaculture, 8: 
307-316. 

PIPE, R. K., 1987, Oogenesis in the marine mus- 
sel Mytilus edulis: an ultrastructural study. 
Marine Biology, 95: 405-414. 

SUN, H., S. KUANG & F. LI, 1996, Studies on 
suitable culture depths and method for scallop 
in Sanggou Bay. Journal of Fisheries Science 
of China, 3: 60-65. 

SUN, J., C. LIN, P. LI, Y. JIN & L. ZHOU, 1997, 
The culture experiment of scallop Chlamys 
farreri in Nanji Islands. Zhejiang College of 
Fisheries, 16: 247-255. 


WHANG, H. J. 8 М. М. KIM, 1973, Study on the 
distribution and ecology of Chlamys farreri 
nipponensis Kuroda around the Taehuksan Is- 
land. Bulletin of National Fisheries Research 
and Development Agency, 11: 25-35 [in Ko- 
геап]. 

WOURMS, J. P., 1987, Oogenesis. Pp. 50-157, 
in: А. С. GIESE, J. $. PEARSE & М. В. PEARSE, eds., 
Reproduction of marine invertebrates, Vol. 9. 
General aspects: seeking unity in diversity. 
Blackwell Scientific Publications, Palo Alto, 
California. xxii + 712 pp. 

YAKOVLEV, У. M. 8 L. $. AFEICHUK, 1995, The 
reproductive cycle of the scallop Chlamys 
farreri in the Sea of Japan. Pp. 193-198, in: P. 
LUBET, J. BARRET & J.-C. DAO, eds., Fisheries, 
biology and aquaulture of pectinids, 8th Inter- 
national Pectinid Workshop, 273 pp. 

YANG, A., Q. WANG, J. KONG, P. LIU, Z. LIU, H. 
SUN, F. LI, R.WANG & M. JIANG, 1999a, Trip- 
loid induction in Chlamys farreri by application 
of 6-dimethyaminopurine. Journal of Fisheries 
of China, 23: 241-247. 

YANG, H., T. ZHANG, J. WANG, P. WANG, У. HE 
& F. ZHANG, 1999b, Growth characteristics of 
Chlamys farreri and its relation with environ- 
mental factors in the intensive raft-culture ar- 
eas of Sishiliwan Bay, Yantai. Journal of 
Shellfish Research, 18: 71-76. 

YOO, J. S., 1976, Korean shells in colour. ligisa, 
Seoul, Korea. 196 pp., 36 pp. pls. [in Korean]. 


Revised ms. accepted 19 January 2008 


MALACOLOGIA, 2008, 50(1-2): 293-302 


DETERMINANTS OF THE DISTRIBUTION OF APPLE SNAILS 
IN HONG KONG TWO DECADES AFTER THEIR INITIAL INVASION 


King-Lun Kwong’, Pak-Ki Wong', Sam S. $. Lau? €. Jain-Wen Qiu” 


ABSTRACT 


This study examined the relative importance of environmental factors and geographic 
isolation on the distribution of apple snails in Hong Kong two decades after their invasion 
from South America. A survey of 61 sites was conducted to collect apple snails and mea- 
sure 18 environmental parameters known to influence mollusk distribution. Identification 
of specimens collected in our study was aided by analysis of DNA sequences, and all 
apple snails collected in Hong Kong were identified as Pomacea canaliculata. Since its 
initial introduction in the early 1980s, the distribution of this invasive snail has only ex- 
panded slightly. Principal component analysis showed that the environmental characteris- 
tics of the study sites varied with habitat. Streams were quite homogenous in chemical 
characteristics and contained little dissolved minerals, whereas ponds, abandoned wet 
farmlands and drainage channels all showed great variations in nutrient loading. Discrimi- 
nant function analysis (DFA) revealed that the inhabited sites typically had high levels of 
phosphate and alkalinity, but the snail was also occasionally found in streams where dis- 
solved ion concentrations and nutrient levels were low. Most of the inhabitable wetlands in 
New Territories have already been occupied by P canaliculata. Because of its unsuitable 
hydrology, Hong Kong Island remains uninhabited by this species. Lantau Island has hab- 


itable sites for this species, and thus is susceptible for invasion in the future. 
Key words: Ampullariidae, Pomacea, hydrology, nutrient, distribution. 


INTRODUCTION 


Apple snails Pomacea spp. (Gastropoda: 
Ampullariidae) are native to the freshwater 
habitats of South America (Martin et al., 2001; 
Cowie et al., 2006). They were first introduced 
into Asia as a delicacy and as a potential food 
crop in the early 1980s, but soon escaped 
aquaculture and established wild populations 
in various countries (Chang, 1985; Mochida, 
1991; Halwart, 1994; Naylor, 1996; Cowie, 
2002; Teo, 2004). Pomacea spp. are omnivo- 
rous, feeding on microorganisms and detritus 
on surface biofilm, eggs and juveniles of other 
snails, animal carrion, and macrophytes, with 
the latter being predominant (Cowie, 2002). 
Their ability to consume macrophytes has 
made them important pests in Asia’s rice farm- 
ing areas and non-agricultural wetlands 
(Carlsson et al., 2004; Carlsson & Lacoursiere, 
2005). 

Few quantitative assessments of the habitat 
of apple snails have been conducted; there- 


fore factors regulating their distribution are 
poorly understood. In its home range of Buenos 
Aires Province, Argentina, high alkalinity has 
been implicated as a possible chemical barrier 
against the expansion of Pomacea canaliculata 
to the western region, whereas mountains likely 
form a geographic barrier against its southern 
expansion (Martin et al., 2001). However, the 
chemical characteristics of the surface water 
in Hong Kong are very different from those in 
Argentina. For instance, mean conductivity in 
its home range (3.5 mS cm”) is 48 to 10 times 
of that in Hong Kong (from 0.07 mS cm" in 
streams to 0.35 mS cm" in flooded furrows; data 
from Yipp, 1990). Thus, the conclusions from 
Martin et al. (2001), especially those regard- 
ing how high alkalinity limiting apple snail dis- 
tribution, may not be applicable to the situation 
in Hong Kong. 

Apple snails invaded Hong Kong in the early 
1980s. The only survey of apple snails was 
conducted in 1988 by Yipp et al. (1991), who 
reported two species — Р maculata and P. 


‘Department of Biology, Hong Kong Baptist University, Hong Kong, P. R. China 
“Environmental Conservation Studies, College of International Education, Hong Kong Baptist University, Hong Kong, 


P. R. China 
“Corresponding author: qiujw@hkbu.edu.hk 


294 KWONG ETAL. 


lineata, both likely misidentifications of Р 
canaliculata — from the vegetable farming ar- 
eas in the northern New Territories bordering 
Guangdong Province, China. They also ana- 
lyzed the water samples from the inhabited 
sites, and found them to have high nutrient 
(nitrogen and phosphorus) contents and con- 
ductivity. But they did not analyze samples 
from the non-inhabited sites, considering that 
these sites remained uncolonized probably 
due to geographical isolation rather than un- 
suitable hydrology. The small land area (1,104 
km?) with various aquatic habitats (i.e., 
streams, drainage channels, ponds, and aban- 
doned wet farmlands) in Hong Kong and the 
length of time (two decades) since the initial 
invasion by apple snails provide an ideal op- 
portunity to examine the relative contribution 
of habitat requirements and geographic barri- 
ers in determining the distribution of this inva- 
sive species. We hypothesize that the current 
distribution pattern of apple snails in Hong 
Kong should reflect their habitat requirements, 
rather than geographic isolation. We surveyed 
various freshwater habitats and determined a 
number of water and sedimentary parameters 
that are known to influence the distribution of 
mollusks (Green, 1971; Lodge et al., 1987; 
Dillon, 2001). Our goals were to update the 
species identity and distribution range of apple 
snails in Hong Kong and to determine envi- 
ronmental parameters that may have influ- 
enced their distribution. The information may 
allow us to predict wetland habitats that are 
suitable for the colonization of apple snails in 
this region, and possibly inform invasions in 
other regions. 


MATERIALS AND METHODS 
Study Area 


Geographically, Hong Kong can be divided 
into four larger areas — New Territories, 
Kowloon, Hong Kong Island and Lantau Island 
— and 261 smaller outlying islands (Fig. 1). 
Kowloon is a highly urbanized area with no 
suitable habitat for aquatic wildlife. New Terri- 
tories can be divided into a relatively flat north- 
ern part and a relatively hilly southern part, 
although there is no distinct boundary between 
the two. Hong Kong Island is connected with 
Kowloon by two underground tunnels, 
whereas Lantau and Kowloon are linked by 
two bridges. Traditionally, agriculture was prac- 
ticed in these three areas, with rice as the 


staple crop. However, due to rapid urbaniza- 
tion, active agriculture can be seen only in 
northern New Territories, with vegetable and 
flower production accounting for about 97% 
of the total value of local crop production 
(AFCD, 2007). 


Sampling and Sample Analysis 


Our preliminary field trips to a number of sites 
showed that, during winter, few apple snails 
reproduced, and the population size was so 
smail that it was difficult to find any individu- 
als. Sampling was thus conducted during sum- 
mer 2006 when apple snail population 
densities were high and their egg clutches 
were easy to see. Sixty-one sites across the 
rural areas of Hong Kong were chosen (Fig. 
1). Some of these were identical to those in 
Yipp et al. (1991). Others were determined by 
checking a local map showing the major fresh- 
water habitats. Overall, there were more sites 
in northern New Territories where there were 
more freshwater habitats. In each site, 
searches for apple snails and their egg 
clutches were performed by the same two 
people (К. L: Kwong & P. К. Wong). These 
were done by checking for up to 250 m of the 
shore of each water body for the presence of 
the colorful egg clutches on emergent vegeta- 
tion or the bank of the site, as well as living 
apple snails among submerged vegetation, on 
the mud surface, or under stones (Martin et 
al., 2001). A site was considered uninhabited 
when no living apple snails or their egg 
clutches were found. When apple snails were 
present, the most morphologically distinct in- 
dividuals were collected, and a selection of 
these individuals were later sent to Robert 
Cowie and Kenneth Hayes of the University 
of Hawaii for species identification using a 
portion of the mitochondrial cytochrome c oxi- 
dase subunit | sequence (COl), as described 
in Cowie et al. (2006). 

Several environmental parameters that are 
known to affect mollusk distribution were de- 
termined. In each site, water depth was deter- 
mined by inserting an extensible fishing rod 
into the middle of the water body until it 
touched the bottom, and measuring the por- 
tion of submerged rod using a tape measure. 
Flow rate was measured with a JDC Flowatch 
meter. Turbidity was measured with an Orion 
AQ4500 turbidometer. Dissolved oxygen (DO), 
pH, and conductivity were measured with an 
Orion 1230 multiparameter meter. Two water 
samples (1L) were taken with P.V.C. bottles 


DETERMINANTS OF APPLE SNAIL DISTRIBUTION 295 


114°00°E 


22°30'N 


22°20’ 


Tsing Yi 


2220 


114%0' 114°20' 


FIG. 1. Map of Hong Kong showing the sampling sites, their habitat type (O — stream, У — drainage 
channel, A — pond, 0 — abandoned wet farmland) and the presence (filled symbol) or absence (open 
symbol) of Pomacea canaliculata. Inset: Amap of China showing the location of Hong Kong. 


and stored at 4°C in a portable cooler. Upon 
return to the laboratory, water samples were 
filtered through a 0.45 um Gelman 66191 
membrane filter. Subsamples were kept at 
appropriate temperature with respect to the 
chemical parameter to be analyzed. All chemi- 
cal parameters, except nitrate and nitrite, were 
measured using standard methods (Eaton et 
al., 2005). Total alkalinity was determined by 
titration with sulphuric acid (Method 2320 B). 
Several captions were measured with a Varian 
Spectr AA-20 atomic absorption spectrometer 
(Method 3111 В for Mg**, K*, Na* and Ее?* and 
Method 3111 D for Ca”*). Total phosphate was 
measured with the ascorbic acid method 
(Method 4500P E) and ammonia with the 
phenate method (Method 4500NH, F). Sul- 
phate content was measured with the turbidi- 
metric method (Method 4500-SO,* E). Nitrate 
and nitrite were measured with Hach low range 
nitrate (Model NI-14, Cat No. 14161-00) and 
nitrite (Model NI-15, Cat No. 21820-00) test 
kits. In sites with a soft substratum, two sur- 
face (0-5 cm) sediment samples (approxi- 
mately 0.5 kg) were collected for analysis of 
the particle size and organic matter. Sub- 
samples were treated with hydrogen peroxide 
and sodium pyrophosphate, wet-sieved with 


an Octagon digital shaker into six graded frac- 
tions (2 mm to < 0.063 mm) and oven-dried 
(50°C) to constant weight. Mean particle size 
and inclusive graphic standard deviation were 
then determined. The remaining samples were 
ignited at 550°C to constant weight to deter- 
mine the content of organic matter. 


Data Analysis 


For each site we calculated the mean value 
of each chemical and physical parameter to 
be used in statistical analysis. Sedimentary 
properties were not used because the data 
were not available from a number of sites 
where the bottom was concrete or rock. Tur- 
bidity data were not included because they 
were affected by instantaneous human distur- 
bance, such as drainage of vegetable gardens, 
in some of the survey sites. Conductivity data 
were not included because they were a com- 
bined measure of inorganic ions. Ammonia, 
nitrite and nitrate values were pooled to give 
an overall estimate of total inorganic nitrogen. 
Except for pH, the data were subjected to log 
(x + 1) transformation. All statistical analyses 
were conducted using SPSS ver. 11 for Win- 
dows (SPSS, Inc.). Principal components 


296 KWONG ETAL. 


TABLE 1. The incidence of Pomacea canaliculata in four types of habitats surveyed, and the corre- 
sponding environmental parameters of each type of habitat. Each datum is mean (Standard devia- 


tion) of all sites of each habitat type. 


Drainage Abandoned 
All sites Stream channel Pond wet farmland 
(n= 61) (n= 8) (n = 38) (eee) (me7) 
No. sites with P. canaliculata 34 2 22 6 4 
% sites with P. canaliculata 95.7 25.0 57.9 75.0 57.1 
Depth (cm) 33.9 29.4 27.0 87.5 15.0 
| (33.7) (14.5) (12.0) (69.2) (6.5) 
OH 6.7 6.4 6.9 6.4 6.5 
(0.7) (0.4) (0.6) (0.8) (0.4) 
DO (mg Г") 4.3 5.7 4.4 3.4 3.5 
(2.0) (2.0) (1.7) (2.6) (2.5) 
Turbidity (NTU) 10.8 5.0 11.2 8.1 18.2 
(17.0) (5.0) (19.9) (7.0) (15.3) 
Flow rate (cm s”) 9.4 13.1 127 0.0 0.0 
(11.9) (11.6) (13.3) (0.0) (0.0) 
Alkalinity (mg CaCO; I") 41.9 9.8 41.2 58.9 63.1 
(41.6) (7.6) (39.5) (56.9) (39.0) 
Calcium (mg I”) 23.8 11.0 23.9 33.2 27.7 
(16.4) (2.6) (16.0) (21.3) (14.6) 
Magnesium (mg Г") 8.1 6.2 7.1 15.9 6.4 
(7.7) (1.4) (2.5) (19.5) (3.2) 
Potassium (mg I”) 57 3.3 5.2 9.7 6.6 
(4.8) (0.6) (2.6) (10.5) (5.1) 
Sodium (mg I”) 54.3 45.2 49.7 91.6 47.5 
(41.7) (12.2) (17.8) (103.9) (22.9) 
Iron (mg I”) 0.7 0.3 0.6 0.4 A 
(1.0) (0.4) (0.8) (0.1) (2.1) 
Sulphate (mg I”) 17.4 5.7 15.2 30.8 24.8 
(25.9) (5.2) (22.7) (34.4) (40.2) 
Inorganic nitrogen (mg |”) 12 0.2 1.5 Re 0.4 
(2.3) (0.5) (2.8) (1.8) (0.3) 
Total phosphate (mg I”) 0.9 0.1 1.0 1.0 A 
(1.4) (0.2) (1.3) Guy (2.5) 
Conductivity (US cm”) 169.2 49.3 172.0 239.3 2415 
(130.0) (15.9) (113.8) 731) (161.1) 
Mean particle size (mm) 03 0.5 0,4 0.1 0.1 
(0.3) (0.2) (0.3) (0.1) (0.1) 
Sorting coefficient 1.9 15 1.8 2.4 1.9 
(0.6) (0.1) (0.6) (1.1) (0.6) 
% organic matter ST 126 33 7.4 5.4 
(3.9) (0.8) (4.1) (3.8) (2.9) 


area. To simplify the data structure and thus 
aid in the interpretation of the results, the com- 
ponent loadings were rotated using the 


analysis (PCA) was performed on the correla- 
tion matrix to characterize the chemical and 
physical characteristics of the water in study 


DETERMINANTS OF APPLE SNAIL DISTRIBUTION 


TABLE 2. Results of principal components analysis of water quality 
parameters (n = 59) showing Varimax rotated component loadings, 
eigen values, and percentage of variance explained. 


Variable PC 1 
Depth -0.328 
pH 0.488 
DO -0.376 
Flow rate -0.282 
Alkalinity 0.792 
Calcium 0.919 
Magnesium 0.558 
Potassium 0.825 
Sodium 0.414 
Iron 0.393 
Inorganic nitrogen 0.545 
Total phosphate 0.601 
Sulphate 0.768 
Eigen value 4.61 
Cumulative % variation 35.4 


Varimax method (NoruSis, 2005). Stepwise 
discriminant function analysis (DFA) was con- 
ducted to determine the major hydrological 
parameters in distinguishing sites that were 
inhabited or uninhabited by apple snails. Pa- 
rameters were entered until no additional pa- 
rameter would significantly increase the 
discriminating power of the function. To deter- 
mine whether the absence of apple snails in 
Lantau and Hong Kong Islands was due to 
unsuitable chemical and physical character- 
istics of the water, we used data from the New 
Territories and Tsing Yi to build up a discrimi- 
nant function, and data from Lantau and Hong 
Kong Islands to cross-validate the result. 


RESULTS 
Species Identity and Distribution of Apple Snails 


Morphologically, the collected apple snails 
showed two color variants — one with a brown 
shell and brown head-foot, and another with a 
yellow shell and yellow head-foot. The brown 
variant was dominant, accounting for over 95% 
of the total individuals found. The yellow vari- 
ant was rarely found. Based on mitochondrial 
COI gene sequences of the 16 most morpho- 
logically distinct specimens, R. Cowie and K. 
Hayes determined our samples to be P. 
canaliculata. Overall, apple snails were found 
in 34 of the 61 surveyed sites (Table 1). They 


Pe POS PC 4 
08351 -0.381 0.495 
-0.071 0.732 0:27 
-0.118 0773 0.060 
0.470 0.117 0395 
-0.340 -0.048 0.254 
-0.114 0.077 0.024 
0.665 -0.108 -0.272 
-0.027 0.057 0.080 
0771 -0.049 -0.396 
-0.544 -0.289 -0.086 
0.326 -0.042 0.564 
-0.002 -0.016 -0.006 
0.091 -0.013 -0.048 

1.94 1.41 1.05 

50.3 Olle 69.3 


297 


were very common in northern New Territo- 
ries, sporadically distributed in southern New 
Territories and Tsing Yi Island, and not present 
in Lantau or Hong Kong Islands (Fig. 1). The 
surveyed sites could be divided into streams, 
drainage channels, ponds and abandoned wet 
farmlands. Apple snails were found in all of 
the four habitat types, but their incidence dif- 
fered, ranging from 25% in streams to 75% in 
ponds. 


Environmental Characteristics of Surveyed Sites 


The environmental parameters differed sub- 
stantially among the four types of habitats 
(Table 1). In general, the streams were char- 
acterized by coarse sediment, high DO, and 
flow rate, but low turbidity, conductivity, alka- 
linity, positive ions and nutrients. The ponds 
usually had high alkalinity, conductivity, cal- 
cium, magnesium, sodium and sulphate. The 
water quality values of the drainage channels 
typically lay between those of the streams and 
ponds. The abandoned wet farmlands were 
similar to the ponds in a number of water qual- 
ity parameters, with notable exceptions of shal- 
lowness and extremely low levels of DO in a 
number of sites. 

Principal components analysis reduced the 
variability of the original 13 parameters into 
four components that had an eigen value 
greater than 1, which in total captured 69.3% 
of the total variability (Table 2). The analysis 


298 KWONG ET AL. 


FIG. 2. À 2-D plot of the principal component loadings based on 13 envi- 
ronmental parameters of 59 surveyed sites. Two of the 61 sites were 
excluded because one had extremely high level of sodium, and another 
had extremely low levels of dissolved minerals. Including them would 
have distorted the distribution pattern of other sites along PC 2. Different 
symbols were used to show the four habitat types (O - stream, У — drain- 
age channel, A — pond, [1 - abandoned wet farmland) and the presence 
(filled symbol) or absence (open symbol) of Pomacea canaliculata. PC 1 
mainly represents nutrient loading and water hardness whereas PC 2 is 
largely dominated by sodium and magnesium. 


resulted in a Kaiser-Meyer-Oklin value of 
0.713, indicating its adequacy in representing 
the variability of the original data. The scores 
of the first two components were presented 
as a 2-D plot (Fig. 2). Along the PC 1 most 
parameters have a positive loading, especially 
calcium, potassium, alkalinity, and sulphate. 
Dissolved oxygen, depth and flow rate had a 
weak and opposite trend (Fig. 2, Table 2). The 
PC 2 was largely dominated by sodium and 
magnesium, both have a positive loading, 
while iron and alkalinity had a weak opposite 
trend. Figure 2 also illustrates the relationship 
between habitat type and hydrology. The 
stream sites are clustered, with negative load- 
ings on PC 1 and low loadings on PC 2, indi- 
cating low nutrient loading in these sites and 
high similarities in chemical and physical char- 
acteristics of the water among them. The pond 
sites are spread widely across PC 1, indicat- 
ing high variation in nutrient loading among 


them, but in general have a positive loading 
on PC 2, reflecting higher salinity, magnesium 
concentrations and depth. The abandoned wet 
farmlands are also spread widely across PC 
1, but on PC 2 they typically have low values. 
Most of the drainage channel sites lay between 
the pond and wet farmland sites on PC 2. Dis- 
solved oxygen and pH were the major deter- 
minants of PC 3, and inorganic nitrogen and 
depth were the major determinant of PC 4. 
These two components, however, only ex- 
plained 19% of the total variability. 


Distribution of Apple Snails in Relation to Wa- 
ter Quality 


The stepwise DFA of chemical and physical 
parameters of the New Territories and Tsing 
Yi sites resulted in total phosphate and alka- 
linity being selected as the parameters to dis- 
criminate between sites with and without apple 


DETERMINANTS OF APPLE SNAIL DISTRIBUTION 299 


snails. The correlation between canonical dis- 
criminate function and total phosphate and 
alkalinity was 0.798 and 0.723, respectively. 
The results (canonical correlation = 0.501, 
Wilks’ À = 0.749, y? = 12.692, P = 0.002) 
showed that, of the 47 sites used, DFA was 
able to correctly classify 74% of the cases. 
The percentage of correctly classified cases 
was higher for the prediction of their absence 
(80%) than that for their presence (69%). High 
total phosphate and alkalinity levels charac- 
terized sites that were inhabited by P. 
canaliculata, whereas low levels of the two 
parameters characterized the uninhabited 
sites. When the discriminant function was ap- 
plied to cross-validate the 12 sites from Hong 
Kong and Lantau Islands, ten sites were cor- 
rectly classified as uninhabited, while two sites 
were misclassified as inhabited. 


DISCUSSION 
Environmental Characteristics of Surveyed Sites 


The surveyed wetland habitats were broadly 
categorized into streams, drainage channels, 
ponds and abandoned wet farmlands (i.e., rice 
paddies or vegetable farms), representing the 
typical freshwater wetland habitats in Hong 
Kong. A comparison showed substantial dif- 
ferences in chemical and physical parameters 
among these different habitats, as well as 
within the same type of habitat, except for 
streams, where the hydrology was uniform 
(Fig. 1, Table 1). Such a situation can be ex- 
plained by the facts that igneous rock under- 
lying most parts of Hong Kong is relatively 
homogenous in chemical composition and 
streams are relatively unaffected by human 
activities (Dudgeon & Corlett, 2004); the ponds 
and drainage channels are affected by nutri- 
ent enrichment from vegetable and flower pro- 
duction or poultry and pig farming, as well as 
periodic flooding during the summer monsoon. 
Some abandoned wet farmlands are anoxic 
because of the high organic content in the soil 
and poor drainage. In general, PCA results on 
hydrological parameters (Fig. 2, Table 2) can 
be explained by the relative intensity of hu- 
man activities and distance from the coast. PC 
1 is an assessment of eutrophication. PC 2 is 
a measure of salinity and magnesium. Over- 
all, the 2-D PCA plot gives a good visual pre- 
sentation of the chemical and physical 
characteristics of the study area (Fig 2). 


Species Identity and Distribution of Apple Snails 


Yipp et al. (1991) reported the presence of 
two species, Ampullaria gigas (Spix, 1827) and 
Ampullaria levior (С. В. Sowerby Ш, 1909). 
According to Cowie (1997), Ampullaria is a 
junior synonym of Pomacea. According to 
Alderson (1925), P. levior is a junior synonym 
of P. lineata, and P. gigas is a junior synonym 
of P. maculata. The molecular results of Cowie 
et al. (2006) showed that only four Pomacea 
species have been introduced into Asia: P. 
canaliculata and P. insularum to many Asian 
countries, P. diffusa to Sri Lanka, and P. 
scalaris to Taiwan. Pomacea lineata is very 
similar to P. canaliculata in shell morphology, 
body color, and egg clutch (Cazzaniga, 2002). 
Molecular sequencing of some of our selected 
specimens showing the greatest morphologi- 
cal variations by R. Cowie and K. Hayes clearly 
revealed the identity of the specimens through- 
out Hong Kong to be Р canaliculata. It is there- 
fore very likely that the apple snail reported 
as A. levior by Yipp et al. (1991) and as P 
lineata by Lam (1994) and Dudgeon & Corlett 
(2004) from Hong Kong is P. canaliculata. 
Pomacea maculata, reported by Yipp et al. 
(1991) as P. gigas, might be a misidentification 
of P. canaliculata that had a larger and thinner 
shell when compared with normal sized indi- 
viduals. In fact, a subspecies of P. canaliculata 
with these conchological characteristics was 
described by Hylton-Scott (1948) from a 
CaCO, poor environment. But it is now well- 
known that P. canaliculata exhibits plasticity 
in shell thickness and shape (Cazzaniga, 
2002; Estebenet & Martin, 2003; Cowie et al., 
2006). 

In the first survey of apple snails in Hong 
Kong conducted in 1988 (Yip et al., 1991), 
shortly after their introduction, they were con- 
fined to the northern New Territories. In this 
survey, we confirmed that northern New Terri- 
tories is still their center of distribution (Fig. 
1). As pointed out by Yipp et al. (1991), the 
frequent occurrence of apple snails in the 
northern New Territories was probably due to 
the relatively flat terrain with an extensive net- 
work of drainage channels plus seasonal mon- 
soons, which must have facilitated its 
dispersion. In southern New Territories and 
Tsing Yilsland, where Yipp and her colleagues 
did not find apple snails, we only recorded iso- 
lated Р canaliculata populations. Such a pat- 
tern indicates a slow expansion of distribution 
range by the apple snails, given that it has 


300 KWONG ET AL. 


been two decades since its initial introduction 
and the total area of New Territories is only 
796 km. 


Can We Predict the Suitability of a Site for 
Apple Snails Based on Water Characteristics? 


Factors affecting the distribution of freshwa- 
ter gastropods can be broadly classified into 
biological (i.e., food, competition, predation), 
chemical and physical (i.e., calcium, salinity, 
flooding, current speed, substratum, tempera- 
ture, desiccation, geographic barrier), or cul- 
tural (i.e., aquarium trade) (Lodge et al., 1987; 
Dillon, 2001; Cowie, 2002). Limiting factors for 
the distribution of Р canaliculata are in gen- 
eral considered to be chemical and physical, 
because there is no evidence to show that 
competition or predation can effectively eradi- 
cate this species (Cowie, 2002; Pizani et al., 
2005; Yusa et al., 2006a). 

Due to the limited number of quantitative 
studies of the habitats of P. canaliculata, there 
is no sufficient information to determine the 
importance of chemical or physical factors in 
regulating its distribution and range expansion. 
Temperature is often a limiting factor for the 
northern distribution of tropical and subtropi- 
cal species, and in fact the abundance of apple 
snail populations in Hong Kong is very low in 
winter months (Cha, 1989). But it is obviously 
not a limiting factor for its distribution in Hong 
Kong, given that the mean air temperature in 
the coldest month (January) is as high as 
15.8°C, and this species can survive through 
the winter in some isolated locations in Japan 
where the monthly temperature can drop to 
8.3°C (Ito, 2002). Although during the winter 
P. canaliculata usually suffer from high mor- 
tality, especially in drained fields, individuals 
living in aquatic environments may acquire 
cold hardiness, and the population may ex- 
plode when water temperature increases in the 
spring (Yusa et al., 2006b; Wada & Matsukura, 
2007). Salinity has been generally considered 
a limiting factor for the distribution of freshwa- 
ter biota. However, it should not be a limiting 
factor for our surveyed sites, because we re- 
corded P. canaliculata in a site with sodium 
concentration reaching as high as 0.35 g I". 
Martin et al. (2001) examined the contribution 
of a number of chemical and physical factors 
to the distribution of P canaliculata in its home 
range in Buenos Aires Province, Argentina. 
They noted that chemical characteristics of the 
water (saline, alkaline waters) formed a physi- 
ological barrier for the expansion of apple 


Snails to the inland, western part of their study 
area, where the mean annual precipitation was 
only as low as 600 mm, whereas mountains 
formed a geographic barrier for its southern 
expansion. 

Our DFA analysis revealed that water of high 
total phosphate and alkalinity was character- 
istic of sites inhabited by P. canaliculata in 
Hong Kong, which is different from the situa- 
tion in its home range. The percentage of cor- 
rectly classified cases for the absence of apple 
snails was high (80%). Of the 15 sites where 
apple snails were absent, only three were 
misclassified. This indicates that chemical and 
physical characteristics of the water can be a 
reasonably good predictor of apple snail ab- 
sence, probably reflecting that most inhabit- 
able sites have already been colonized. The 
three exceptions were all drainage channels, 
where the bottom and walls were lined with 
cement and water velocity can be very high 
during the wet season, a condition discourag- 
ing the establishment of apple snail popula- 
tions. High current velocity also discouraged 
the accumulation of mud, which might be a 
refuge for apple snails. The lower percentage 
of correctly classified cases for the presence 
of apple snails (69%) reflected the fact that 
apple snails were found in ten sites of appar- 
ent unsuitable hydrology. These were mainly 
streams and drainage channels with relatively 
low alkalinity and dissolved mineral concen- 
trations. Although the densities of apple snails 
in these sites were very low, and it is not sure 
whether they were transported by water from 
the surrounding areas when flooding occurred 
during the summer monsoons, their presence 
indicates that they could at least survive in 
these habitats for a period of time. Our obser- 
vation indicated that they are especially prone 
to drift in currents during mating. However, it 
remains to be tested whether apple snails can 
complete their life cycle in such fast flowing 
streams. Dudgeon & Corlett (2004) reported 
that, of the other common species of freshwa- 
ter gastropods in Hong Kong, only Brotia 
hainanensis, which has a strong foot to hold 
on to rocks in stream beds, can reproduce to 
reach substantial densities in such habitats. 
Apple snails grown in tap water with a Ca** 
concentration of 11 mg/L last year in our lab 
from neonates had thin shells. When the wa- 
ter was supplemented with a small bag of coral 
fragments, this shell thinning phenomenon 
disappeared. From the field survey, the low- 
est Ca** concentration in the sites where apple 
snails were present was 9.1 mg/L. These ob- 


DETERMINANTS OF APPLE SNAIL DISTRIBUTION 301 


servations indicate the potential limiting effect 
of calcium and alkalinity, but more data from 
laboratory controlled experiments and field 
transplant experiments are required to deter- 
mine whether the chemical characteristics of 
the stream water or flow rate is the limiting 
factor for the distribution of Р canaliculata. 

Of particular interest was that two sites on 
Lantau Island that were classified as inhabit- 
able, yet lacked apple snails (Fig. 1). Of these 
two sites, Pui O is an abandoned paddy field, 
and Tong Fuk is a cement-lined drainage 
channel. The gastropods Melanoides tuber- 
culata and Physella acuta where found in Pui 
O and Tong Fuk, respectively. Elsewhere in 
the surveyed area throughout New Territories, 
these gastropods were often found together 
with Р canaliculata. Such a sympatric distri- 
bution pattern indicates that apple snails may 
share some common habitat requirements 
with these two freshwater snails, or that they 
have the same dispersal mechanism (i.e., 
transported by water in streams and chan- 
nels). Additionally, the input of nutrients from 
animals or domestic waste water and other 
salts associated past agricultural activities, 
such as calcium in lime, may be the reason 
for these sites being classified as inhabitable 
by apple snails in the DFA. Because our data 
provided evidence that chemical characteris- 
tics of the water in these two sites was suit- 
able for the colonization of apple snails, their 
absence indicates that they have not been 
introduced into these sites due to geographic 
isolation. 


Implications for Management 


Although P. canaliculata preferred waters of 
high levels of alkalinity and nutrients charac- 
teristic of vegetable farming areas in northern 
New Territories, it was still sometimes found 
in streams with low levels of alkalinity and 
nutrients. The unsuitable chemical character- 
istics of the water on Hong Kong Island would 
prevent apple snails from establishing a popu- 
lation on this island. However, the presence 
of two sites of favorable chemical character- 
istics of the water on Lantau Island indicates 
a possible risk of future colonization by apple 
snails on this island. It is important to note that, 
once introduced to favorable habitats, apple 
snails may spread very quickly, and it is very 
difficult to eradicate them. 

Throughout the study we have emphasized 
the importance of environmental characteris- 
tics that may affect the colonization of apple 


snails. A recent study has shown that, once 
colonization has taken place, apple snails can 
affect the habitat, such as decreasing the bio- 
mass of wetland macrophytes and increasing 
the nutrient contents of the water (Carlsson et 
al., 2004; Carlsson & Lacoursiére, 2005). It 
can not be excluded that the presence of apple 
snails may also contribute to water quality and 
the patterns of PCA and DFA observed in this 
study. These recent results should also be 
considered in the management of apple snails. 
No matter whether the chemical characteris- 
tics of the water are apparently unsuitable for 
apple snails, caution should be taken by the 
government to prevent the apple snails from 
spreading to the wetlands of Hong Kong and 
Lantau Islands where there are macrophytes. 
This is particularly relevant because a num- 
ber of wetlands on these islands are being 
marketed as ecotourism attractions. 


ACKNOWLEDGMENTS 


We thank Robert Cowie and Ken Hayes (Uni- 
versity of Hawaii) for identifying the snails, 
David Dudgeon (University of Hong Kong) and 
Takashi Wada (National Agricultural Research 
Center, Kyushu Okinawa Region, Japan) for 
providing some useful references, Ricky N. S. 
Wong for helpful comments on the manuscript, 
and three reviewers for critical comments. This 
study was supported by a grant to JWQ from 
Environment and Conservation Fund, Hong 
Kong. 


LITERATURE CITED 


AFCD [Fisheries and Conservation Department, 
Hong Kong Special Administrative Region], 
2007, Agriculture in Hong Kong. http://www. 
afcd.gov.hk/english/ agriculture/agr_hk/ 
agr_hk.html [Cited 25 June 2007]. 

ALDERSON, E. G., 1925, Studies in Ampullaria. 
Cambridge, W. Heffer & Sons. xx + 102 pp. 

CARLSSON, М. O. L., С. BRONMARK & L.-A. 
HANSSON, 2004, Invading herbivory: the 
golden apple snail alters ecosystem function- 
ing in Asian wetlands. Ecology, 85: 1575-1580. 

CARLSSON, N. O. L. & J. O. LACOURSIERE, 
2005, Herbivory on aquatic vascular plants by 
the introduced golden apple snail (Pomacea 
canaliculata) in Lao PDR. Biological Invasions, 
7: 233-241. 

CAZZANIGA, N. J., 2002, Old species and new 
concepts in the taxonomy of Pomacea (Gas- 
tropoda: Ampullariidae). Biocell, 26: 71-81. 

CHA, M. W., 1989, Interspecific interactions 
among freshwater Gastropoda in Hong Kong. 


302 KWONG ET AL. 


Unpublished M. Phil. Thesis, Hong Kong Poly- 
technic, Hong Kong. 

CHANG, W. C., 1985, The ecological studies on 
the Ampullaria snails. Bulletin of Malacology, 
Republic of China, 11: 43-51. 

COWIE, БК. H., 1997, Pila Röding, 1798 and 
Pomacea Perry, 1810 (Mollusca, Gastropoda): 
proposed placement on the Official List, and 
Ampullariidae Gray, 1824: proposed confirma- 
tion as the nomenclaturally valid synonym of 
Pilidae Preston, 1915. Bulletin of Zoological 
Nomenclature, 54: 83-88. 

COWIE, В. H., 2002, Apple snails (Ampullariidae) 
as agricultural pests: their biology, impacts and 
management. Pp. 145-192, in: С. М. BAKER, ed., 
Molluscs as crop pests. Wallingford, New 
Zealand, CABI Publishing. xii + 468 pp. 

COWIE, К. H., К. A. HAYES & S. С. THIENGO, 
2006, What are apple snails? Confused tax- 
onomy and some preliminary resolution. Pp. 
3-23, in: R. С. JOSHI, ed., Global advances in 
ecology and management of golden apple 
snails. Muñoz, Nueva Ecija, Philippine Rice 
Research Institute. x + 588 pp. 

DILLON, R. T. Jr., 2000, The ecology of fresh- 
water molluscs. Cambridge, Cambridge Uni- 
versity Press. xii + 509 pp. 

DUDGEON, D. & R. CORLETT, 2004, The ecol- 
ogy and biodiversity of Hong Kong. Hong Kong, 
Friends of the Country Parks / Joint Publish- 
ing. 336 pp. 

EATON, A. D., L. S. CLESCERI, Е. W. RICE, A. 
E. GREENBERG, eds., 2005, Standard meth- 
ods for the examination of water and waste- 
water, 21% ed. Washington, D.C., American 
Public Health Association. Various pagination. 

ESTEBENET, А. L. & Р. В. MARTIN, 2003, Shell 
interpopulation variation and its Origin in 
Pomacea canaliculata (Gastropoda: Am- 
pullariidae) from Southern Pampas, Argentina. 
Journal of Molluscan Studies, 69: 301-310. 

GREEN, R. H., 1971, A multivariate statistical 
approach to the Hutchinsonian niche: bivalve 
mollusks of central Canada. Ecology, 52: 543- 
560: 

HALWART, M., 1994, The golden apple snail 
Pomacea canaliculata in Asian rice farming 
systems: present impact and future threat. /n- 
ternational Journal of Pest Management, 40: 
199-206. 

HYLTON-SCOTT, М. I., 1948, Moluscos del 
noroeste Argentino. Acta Zoologica Lilloana, 
6: 241-274. 

ITO, K., 2002, Environmental factors influencing 
overwintering success of the golden apple 
snail, Pomacea canaliculata (Gastropoda: 
Ampullariidae), in the northern most popula- 
tion in Japan. Applied Entomology and Zool- 
ogy, 37: 655-661. 

LAM, P. K. S., 1994, Intraspecific life-history 
variation in Radix ‘plicatulus (Gastropoda: 


Pulmonata: Lymnaeidae). Journal of Zoology, 
London, 232: 435-446. 

LODGE, D. M., К. BROWN, $. KLOSIEWSKI, 
R. STEIN, ay COVICH, B. LEATHERS & es 
BRÓNMARK, 1987, Distribution of freshwater 
snails: spatial scale and relative importance of 
physiological and biotic factors. American Ma- 
lacological Bulletin, 5: 73-84. 

MARTIN, P. R., А. |. ESTEBENET & М. J. 
CAZZANIGA, 2001, Factors affecting the dis- 
tribution of Pomacea canaliculata (Gastropoda: 
Ampullariidae) along its southernmost natural 
limit. Malacologia, 43: 13-23. 

MOCHIDA, O., 1991, Spread of freshwater Po- 
macea snail (Pilidae, Mollusca) from Argentina 
to Asia. Micronesica, Supplement, 3: 51-62. 

NORUSIS, M., 2006, SPSS 15.0 Statistical pro- 
cedures companion. Prentice-Hall, Upper 
Saddle River, New Jersey. xxii + 626 pp. 

NAYLOR, R., 1996, Invasions in agriculture: As- 
sessing the cost of the golden apple snail in 
Asia. Ambio, 25: 443-448. 

PIZANI, М. М. a L. ESTEBENET 8 P. В. MARTIIN, 
2005, Effects of submersion and aerial ехро- 
sure оп clutches and hatchlings of Pomacea 
canaliculata (Gastropoda: Ampullariidae). 
American Malacological Bulletin, 20: 55-63. 

TEO, S. S., 2004, Biology of the golden apple 
snail, Pomacea canaliculata (Lamarck, 1822), 
with emphasis on responses to certain envi- 
ronmental conditions in Sabah, Malaysia. Mol- 
luscan Research, 24: 139-148. 

WADA, T. & K. MATSUKURA, 2007, Seasonal 
changes in cold hardiness of the invasive fresh- 
water apple snail, Pomacea canaliculata 
(Lamarck) (Gastropoda: Ampullariidae). 
Malacologia, 49: 383-392. 

YIPP, M. W., 1990, Distribution of the schisto- 
some vector snail, Biomphalaria straminea 
(Pulmonata: Planorbidae) in Hong Kong. Jour- 
nal of Molluscan Studies, 56: 47-55. 

YIPP, М. W., M. W. СНА & X. Y. LIANG, 1991, A 
preliminary impact assessment of the introduc- 
tion of two species of Ampullaria (Gastropoda: 
Ampullariidae) into Hong Kong. Pp. 393-397, 
in: C. MEIER-BROOK, ed., Proceedings of the 
Tenth International Malacological Congress, 
Tubingen 1989, Baja, Hungary, UNITAS 
Malacologia, 2 vols. vi + 636 pp. 

YUSA, Y., N. SUGIURA & T. WADA, 2006a, 
Predatory potential of freshwater animals on 
an invasive agricultural pest, the apple snail 
Pomacea canaliculata (Gastropoda: Am- 
pullariidae), in southern Japan. Biological In- 
vasions, 8: 137-147. 

YUSA, Y., Т. WADA & $. TAKAHASHI, 2006b, 
Effects of dormant duration, body size, self- 
burial and water condition on the long-term 
survival of the apple snail, Pomacea cana- 
liculata (Gastropoda: Ampullariidae). Applied 
Entomology and Zoology, 41: 627-632. 


Revised ms. accepted 19 February 2008 


MALACOLOGIA, 2008, 50(1-2): 303-318 


EVOLUTIONARY RELATIONSHIPS OF POPENAIAS POPEII AND 
THE EARLY EVOLUTION OF LAMPSILINE BIVALVES (UNIONIDAE): 
PHYLOGENETIC ANALYSES OF DNAAND AMINO ACID SEQUENCES 
FROM F AND M MITOCHONDRIAL GENOMES 


Eric С. Chapman’, Mark E. Gordon‘, Jennifer M. Walker, Brian К. Lang’, David С. 


Campbell, С. Thomas Watters®, Jason P. Curole®, Helen Piontkivska' & Walter R. Hoeh*** 


ABSTRACT 


Lampsiline bivalves typically are considered a tribe within the Unionidae (Ambleminae: 
Lampsilini), and they display extraordinary morphological adaptations for reproduction. 
Recent studies have weakly corroborated the monophyly of the Lampsilini, but evolution- 
ary relationships within the tribe and its sister lineage have yet to be elucidated convinc- 
ingly. However, these determinations are necessary to better understand the evolution of 
the spectacular morphological diversity present in lampsilines, as well as the specific cir- 
cumstances surrounding the group's origin. To clarify these matters, phylogenetic analy- 
ses were carried out on 2,310 nucleotide and 770 amino acid position matrices containing 
sequences from five protein coding gene regions on the F and M mitochondrial genomes 
from 21 amblemine species. Nodal support values on the best Bayesian inference tree 
robustly confirm the monophyly of lampsilines and a clade containing the following well- 
supported relationships: (((lampsilines, Popenaias) Plectomerus) Amblema). Furthermore, 
a maximum likelihood estimate of ancestral character states indicates that the ectobranchy 
observed in lampsilines + Popenaias is homologous and was derived from a tetragenous 
ancestral lineage. The sister taxon status of P. popeii to the traditional lampsiline taxa and 
the occasional use of the inner demibranchs for brooding suggest that this species could 
still retain character states of the lampsiline ancestral lineage. Therefore, additional stud- 
ies of morphology, reproduction, phylogeography and ecology for Popenaias, Amblema, 
Plectomerus, and other taxa within the Amblemini could clarify the circumstances sur- 
rounding the origin of the lampsiline bivalves. 

Key words: Cytochrome с oxidase subunits | & Il (coxT, cox2), DUI, phylogenetics, 
Ambleminae, Unionidae, Popenaias popeii. 


INTRODUCTION 


Lampsiline bivalves (Unionidae: Amblemi- 
nae: Lampsilini; sensu Davis & Fuller, 1981) 
exhibit a considerable array of morphological 
and anatomical adaptations (Zanatta & 
Murphy, 2006). Taxa are endemic to eastern 
North and Central America, with 119 species 
in 21 genera currently recognized in the United 
States (Burch, 1975; Davis & Fuller, 1981; 
Turgeon et al., 1998; Roe & Hartfield, 2005). 
Generally regarded as a monophyletic group 


(Heard & Guckert, 1971; Davis & Fuller, 1981; 
Lydeard et al., 1996; Graf & Ó Foighil, 2000; 
Campbell et al., 2005; Zanatta 8 Murphy, 2006; 
but see Frierson, 1927), it was first recognized 
as such by Ihering (1901); however, diagnos- 
tic criteria were not specified until Ortmann 
(1910). Among these apparent diagnostic char- 
acteristics, the following features have been 
considered particularly definitive: (1) dorsal 
margin of inner lamina of inner gills generally 
entirely connected with abdominal sac (Ort- 
mann, 1912); (2) mantle edge anteroventral to 


‘Department of Biological Sciences, Kent State University, Kent, Ohio 44242, U.S.A. 

2New Mexico Museum of Natural History and Science, 1801 Mountain Road NW, Albuquerque, New Mexico 87104, U.S.A. 
¿New Mexico Department of Game and Fish, 1 Wildlife Way, Santa Fe, New Mexico 87507, U.S.A. 

“Biodiversity and Systematics, Department of Biological Sciences, University of Alabama, Tuscaloosa, Alabama 35847, U.S.A. 
5Ohio Biological Survey and Aquatic Ecology Lab, Ohio State University, 1315 Kinnear Road, Columbus, Ohio 43212, U.S.A. 
“Department of Biological Sciences, University of Southern California, Los Angeles, California 90089, U.S.A. 

*Current address: Department of Biological Sciences, University of Southern Mississippi, Long Beach, Mississippi 39560, U.S.A. 


**Corresponding author: whoeh@kent.edu 


304 CHAPMAN ET AL. 


the incurrent, branchial opening in females 
crenulated, papillose, or with a structural elabo- 
ration, for example, a caruncle or flap whose 
motion is customarily interpreted as a lure to 
attract hosts (Ortmann, 1912; Heard & Guckert, 
1971; Fuller, 1975; Davis & Fuller, 1981); (3) 
when gravid, marsupial water tubes extending 
below the ventral margin of the branchial fila- 
ments (Ortmann, 1912; Fuller, 1975; Davis & 
Fuller, 1981); and (4) larvae (glochidia) re- 
leased from the mother through pores that 
develop at the ventral margin of the marsupial 
water-tubes (Ortmann, 1912). Ortmann, refer- 
ring to the Lampsilinae, stated (1912: 301): 
“The latter physiological character [#4 above] 
is unique, and is found only in this subfamily.” 
However, these “diagnostic” morphological 
characteristics likely do not represent unique 
and unreversed character states for lampsilines 
but rather generalities noted by Ortmann 
(1910, 1912) (e.g., glochidia are not released 
from the ventral margin of the marsupium in 
Cyprogenia, Hamiota, and Dromus — Cham- 
berlain, 1934; Haag et al., 1995; Jones et al., 
2004). Therefore, the answer to the question, 
“What is a lampsiline?” is potentially conten- 
tious. Evolutionary relationships among the 
nominal lampsiline genera are still poorly un- 
derstood. Cyprogenia, Cyrtonaias, Dromus, 
Friersonia, Obliquaria, and Ptychobranchus 
usually are acknowledged as being relatively 
primitive members of the Lampsilini based on 
the marsupial morphology and relative lack of 
conchological sexual dimorphism (Ortmann, 
1912; Heard & Guckert, 1971; Fuller, 1975; 
Davis & Fuller, 1981). The remaining genera 
(“heterogenae” [i.e., posteriorly located mar- 
supia] of Simpson, 1900, 1914; Heard & 
Guckert, 1971; Davis & Fuller, 1981) are re- 
garded as a relatively derived monophyletic 
group. Rather than relying on formal phyloge- 
netic analyses, most assessments regarding 
supra- and inter-generic phylogenetic relation- 
ships within the Lampsilini were based on rela- 
tively small numbers of characteristics and the 
assumption that relatively simple structures 
are ancestral to relatively complex, special- 
ized structures. Recent studies of lampsiline 
evolutionary relationships (Campbell et al., 
2005; Zanatta & Murphy, 2006) have not ro- 
bustly corroborated the monophyly of the taxon 
nor have they convincingly elucidated the re- 
lationships within the group or its sister lineage. 
These determinations are necessary to better 
understand the specific circumstances sur- 
rounding the origin of the Lampsilini and the 


evolution of its extraordinary morphological 
diversity. 

Campbell et al. (2005) and Zanatta & Murphy 
(2006) currently represent the most compre- 
hensive studies of lampsiline bivalve phylog- 
eny. Their results associate Popenaias popeii 
(Lea, 1857) either (1) with a clade containing 
Amblema (Campbell et al., 2005: figs. 1, 2; 
Zanatta & Murphy, 2006: fig. 1) or (2) as a po- 
tential sister taxon to the Lampsilini (Zanatta 
& Murphy, 2006: fig. 2). Although Bayesian in- 
ference (BI), posterior probability (PP) indicat- 
ing amblemine affinities for Popenaias was 
quite high in Campbell et al. (2005: 0.99, fig. 
2), the PP of the BI analysis in Zanatta & 
Murphy (2006: fig. 2) and maximum parsimony 
(MP) bootstrap percentages in both studies 
were less than 0.5 and 55, respectively. Origi- 
nally described as Unio popeii Lea, 1857, pre- 
vious Classifications have been rather 
inconsistent regarding the phylogenetic pro- 
pinquity of this species. Simpson (1900, 1914), 
Cockerell (1902), and Hinckley (1907) retained 
it within Unio Philipsson, 1788, but Pilsbry 
(1909a, b) listed it as a Lampsilis Rafinesque, 
1820. Based on anatomical features includ- 
ing a presumed ectobranchus marsupium and 
similarities of anatomy, shell shape, and beak 
sculpture, Ortmann (1912) classified it under 
Elliptio Rafinesque, 1820. Frierson (1927) also 
classified it as an Elliptio but erected a new 
subgenus, Popenaias, with it as the type spe- 
cies and including two Mexican taxa. Heard & 
Guckert (1971) elevated Popenaias to generic 
status with two species (popeii and buckleyi 
Lea, 1843) and proposed Popenaiadinae for 
Popenaias and Cyrtonaias, based primarily on 
reproductive periodicity and “homogenae” 
marsupial morphology. However, Heard (1974) 
abandoned this subfamilial classification after 
determining that the diagnoses for Popenaia- 
dinae were based on species specific charac- 
ters not warranting higher classification. 
Johnson (1972, 1999) re-established E. 
buckleyi and reclassified Popenaias into Pleu- 
robeminae Hannibal, 1912, respectively. Fuller 
(1975) established Cyrtonaias as a lampsiline. 
Subsequently, Smith et al. (2003) concluded 
that P popeii had pleurobemine-like (Amble- 
minae: Pleurobemini) anatomy and, while typi- 
cally exhibiting ectobranchy, demonstrated 
that P popeii can brood embryos in the inner 
demibranchs facultatively, an anatomical con- 
dition distinct from the brooding condition in 
the above-mentioned genera. Given the 
above, the evolutionary relationships of both 


EVOLUTIONARY RELATIONSHIPS OF POPENAIAS POPEII 305 


the Lampsilini and P. popeii remain inad- 
equately understood. 

Freshwater unionoidean bivalves, as well as 
representatives of the two marine bivalve or- 
ders Mytiloida and Veneroida, exhibit doubly 
uniparental inheritance (DUI) of mitochondrial 
DNA (mtDNA), which involves distinct mater- 
nal (F) and paternal (M) transmission routes 
concomitant with highly divergent gender-as- 
sociated mtDNA genomes (Hoeh et al., 1996, 
2002; Liu et al., 1996; Curole & Kocher, 2002, 
2005; Walker et al., 2006; for a general re- 
view of DUI, see Breton et al., 2007). Females 
transmit their mitochondria (carrying F mtDNA) 
to sons and daughters, as in standard mater- 
nal inheritance, but males effectively transmit 
their mitochondria (via sperm carrying M 
mtDNA) only to sons (e.g., Sutherland et al., 
1998; but see Obata et al., 2007; Chakrabarti 
et al., 2007). In males, F mtDNA predominates 
in the somatic tissues while principally M 
mtDNA is found in the testes. Thus, this ge- 
netic system yields homoplasmic (= contain- 
ing a single mtDNA type) female and 
heteroplasmic (= containing multiple mtDNA 
types) male individuals. Intra- and inter-spe- 
cific comparisons suggest that the M genome 
is evolving more rapidly than the F genome 
(Skibinski et al., 1994; Rawson & Hilbish, 
1995; Stewart et al., 1995; Liu et al., 1996; Hoeh 
et al., 2002; Krebs, 2004). The F and M mito- 
chondrial genomes of unionoidean bivalves 
form reciprocally monophyletic groups (Curole 
& Kocher, 2002, 2005; Hoeh et al., 1996, 2002: 
Walker et al., 2006) and are highly divergent 
(Mizi et al., 2005). Fossil evidence suggests 
that the F/M divergence, and concomitant in- 
dependent evolution of F and M mitochondrial 
genomes, occurred > 200 MYA (Watters, 
2001). Phylogenetic analyses of unionoidean 
species, using DNA sequences from both 
mtDNA lineages within a species as distinct 
terminals, typically yields well supported F and 
M clades with very similar topologies (Hoeh 
et al., 1996, 2002; Curole & Kocher, 2002, 
2005; Krebs, 2004; Walker et al., 2006). Thus, 
concatenating F and M sequences (i.e., using 
F and M sequences additively to represent a 
single terminal) for phylogenetic analyses is 
justifiable, and the resulting trees are often 
more robustly supported (e.g., Hoeh et al., 
2002; Walker et al., 2006) than those restricted 
to analyses of only F genome sequences (e.g., 
Campbell et al., 2005; Zanatta & Murphy, 
2006). 

The phylogenetic relationships of Popenaias 
popeli and other amblemine-like species near 


the ancestral lineage of the lampsiline clade 
remain unresolved (e.g., Campbell et al., 2005; 
Zanatta & Murphy, 2006). Inconsistencies of 
morphologically based classifications with the 
phylogenetic trees presented in Campbell et 
al. (2005) and Zanatta & Murphy (2006) fur- 
ther indicate the need for additional assess- 
ment of the evolutionary relationships of these 
taxa. To clarify these matters, matrices con- 
taining 2,310 nucleotide (nt) and 770 amino 
acid (a.a.) positions were constructed with 
sequences generated from five gene regions 
encoded on the F and M mitochondrial ge- 
nomes of 21 amblemine species. Our phylo- 
genetic analyses addressed phylogenetic 
relationships in the Ambleminae with special 
reference to the Lampsilini and pertinent evo- 
lutionary relationships indicated in the evolu- 
tionary trees of Campbell et al. (2005) and 
Zanatta & Murphy (2006). 


MATERIALS AND METHODS 
Taxa Used 


We obtained sequences from a thorough 
cross-section (п = 21 species; Table 1) of the 
unionoidean bivalve subfamily Ambleminae, 
including 14 genera (16 species) represent- 
ing the Amblemini and Lampsilini (sensu 
Campbell et al., 2005), two genera (two spe- 
cies) from the Pleurobemini, one genus (two 
species) representing the Quadrulini, and one 
species from the Gonideini. Collection locality 
information for the specimens utilized herein 
is presented in Appendix |. The use of /n- 
versidens japanensis (Gonideini) as the 
outgroup in our phylogenetic analyses is jus- 
tified by the results of Campbell et al. (2005) 
and Walker et al. (2006). Paleontological evi- 
dence suggests that the sequences analyzed 
herein diverged from a common ancestor 
> 65 MYA (Watters, 2001). 


DNA Sequencing 


Gender of each specimen was determined 
by microscopical examination of gonadal tis- 
sues. Total genomic DNA was isolated from 
mantle and testes using the Qiagen DNeasy 
animal kit. The largely M-specific primer pair 
from Chakrabarti et al. (2006) along with those 
in Walker et al. (2007) were used to amplify 
the Mcox2-cox7 junction region (Curole € 
Kocher, 2002) from testicular tissue-based 
DNA isolates. These primers amplified an 


306 


~1.7 kbp fragment from the M genomes. A 
largely F-specific primer pair (Walker et al., 
2006) was used to amplify the corresponding 
Fcox2-cox1 junction region from mantle tissue- 
based DNA isolates. These primers amplified 
а ~1.1 kbp fragment from the Е genome. The 


CHAPMAN ET AL. 


actual number of nucleotides used in the phy- 
logenetic analyses (2,310 nt) is somewhat less 
that that suggested by the above fragment 
lengths due to the deletion of primer se- 
quences, intergenic spacer regions and regions 
containing indels (see Results and Discussion). 


TABLE 1. Species and GenBank accession numbers for the DNA sequences used in this study. 
Female-transmitted cytochrome с oxidase subunit | (FcoxT); female-transmitted cytochrome с oxi- 
dase subunit Il (Fcox2); male-transmitted cytochrome с oxidase subunit | (McoxT); and male-trans- 
mitted cytochrome с oxidase subunit II (Mcox2). Tribal assignments are as in Campbell et al. (2005: 


fig. 2 and text). 


Species 


Inversidens japanensis 
(Lea, 1859) 
Quadrula quadrula 
(Rafinesque, 1820) 
Quadrula refulgens 
(Lea, 1868) 
Fusconaia flava 
(Rafinesque, 1820) 
Pleurobema sintoxia 
(Rafinesque, 1820) 
Amblema plicata 
(Say, 1817) 
Popenaias popeii 
(Lea, 1857) 
Actinonaias ligamentina 
(Lamarck, 1819) 
Cyrtonaias tampicoensis 
(Lea, 1838) 
Glebula rotundata 
(Lamarck, 1819) 
Hamiota subangulata 
(Lea, 1840) 
Lampsilis hydiana 
(Lea, 1838) 
Lampsilis ovata 
(Say, 1817) 
Lampsilis straminea 
(Conrad, 1834) 
Lemiox rimosus 
(Rafinesque, 1831) 
Obliquaria reflexa 
(Rafinesque, 1820) 
Obovaria olivaria 
(Rafinesque, 1820) 


Plectomerus dombeyanus 


(Valenciennes, 1827) 


Ptychobranchus fasciolaris 


(Rafinesque, 1820) 
Toxolasma lividus 
(Rafinesque, 1831) 


Tribe 


Gonideini 
Quadrulini 


Quadrulini 


Pleurobemini 


Pleurobemini 


Amblemini 
Amblemini 
Lampsilini 
Lampsilini 
Lampsilini 
Lampsilini 
Lampsilini 
Lampsilini 
Lampsilini 
Lampsilini 
Lampsilini 
Lampsilini 
Lampsilini 
Lampsilini 


Lampsilini 


Venustaconcha ellipsiformis Lampsilini 


(Conrad, 1836) 


Fcox1 


AB055625 


EF033268 


EF033269 


EFOSS261 


EF033253 


En035258 


EF033257 


EFOS3Z05 


Er035259 


EF033264 


EF033266 


EF033270 


EF033262 


EF033271 


EFO33256 


EF033254 


EF033207 


EF033252 


EFO33265 


EF033255 


EF033260 


Fcox2 


AB055625 


EF033288 


AF517643 


EF033281 


ER033273 


EF033278 


ЕО 52 77 


EF033283 


EROSS27© 


EF033284 


EF033286 


ÉROSS202 


EF033289 


EF033276 


EF033274 


EF033287 


EE033272 


EF033285 


EFOSS275 


EF033280 


Mcox1 


AB055624 


EF033308 


EF033309 


EE033307 


EF033291 


EE033295 


EF033294 


EFO33300 


EFOS5299 


EFO33304 


ЕР 5505 


EFRUSS296 


EFOS33903 


EFOSS297 


ER083302 


ER033292 


EFO33306 


EFOS3290 


EFOS3301 


ER033293 


EF033296 


Mcox2 
AB055624 


EF033328 


AF517638 


21093327 


EROSS3 


ЕВ SS 


EF033314 


EF033320 


Er 033319 


EF033324 


ЕР 


Во 


ÉPOUSSSES 


ÉFOSSS 17 


ÉF053322 


EROSSSIZ 


EF033326 


EF033310 


ER0338321 


EF033313 


EF033316 


EVOLUTIONARY RELATIONSHIPS OF POPENAIAS РОРЕП 307 


PCR reactions consisted of 1X Qiagen PCR 
buffer, 0.2 mM each dNTP, 0.5 M each primer, 
1U Qiagen Tag and ~20 ng of template DNA. 
Reactions using the M-specific primers were 
cycled at 94°C for 60 s, 50°C for 60 s, and 
72°C for 120 s for a total of 40 cycles. Reac- 
tions involving the F-specific primers followed 
the same profile as above, but were annealed 
at 46°C. The above PCR primers ultimately 
yielded F and Mcox2-cox1 DNA sequences 
obtained via cycle sequencing with Perkin 
Elmer AmpliCycle Sequencing Kits. Sequenc- 
ing primers were identical in sequence to the 
PCR primers and sequencing template purifi- 
cation was done following Folmer et al. (1994). 
Sequences were visualized using Li-Cor 
4200L-2 and 4200S-2 DNA sequencers. For- 
ward and reverse sequencing reads were as- 
sembled and verified using AlignIR (version 2.0, 
LI-COR, Inc.), and final sequence alignments 
were completed manually with MacClade ver- 
sion 4.0 (Maddison & Maddison, 2000). 


Phylogenetic Analyses 


Our Mcox2 sequences were aligned with 
unionoidean Fcox2 nucleotide sequences 
(which have a uniform length) obtained from 
the GenBank and herein to determine the 
boundaries and length of Mcox2e, the 
hypervariable coding extension of the Mcox2 
gene specific to unionoid M mtDNA (Curole & 
Kocher, 2002; Walker et al., 2007). The 5’ end 
of Mcox2e is designated as the nucleotide of 
the Mcox2 sequence that aligns with position 
one in the stop codon of the Fcox2 sequences. 
The 3’ end of Mcox2e is the stop codon for 
Mcox2. The F and Mcox2-cox1 nucleotide 
sequences were translated to protein se- 
quences using the Drosophila mtDNA genetic 
code. All DNA sequences generated herein 
were submitted to the GenBank database 
(Table 1). 

Phylogenetic trees were estimated using 
Bayesian inference (Bl), neighbor-joining (NJ) 
and maximum parsimony (MP) approaches. 
Bayesian analyses were conducted using the 
program MrBayes (version 3.1.2: Huelsenbeck 
& Ronquist, 2005; Ronquist & Huelsenbeck, 
2003). For nucleotide sequences, Bayesian 
searches were run for 10 million generations 
with 10 search chains, and the data were par- 
titioned by gene region and codon position (five 
gene regions x three codon positions for the 
full data-set; four gene regions x three codon 
positions for the data-set with the Mcox2e se- 
quences removed), saving 10,000 trees (one 


tree saved every 1,000 generations) and us- 
ing GTR + С + | substitution model (Rodriguez 
et al., 1990) as selected by the program 
Modeltest (Posada & Crandall, 1998). To al- 
low each partition to have its own set of pa- 
rameter estimates, revmat, tratio, statefreg, 
shape, and pinvar were all unlinked during the 
analysis. Burn-in was determined by visual in- 
spection of the likelihood score plots obtained 
as the trees were written to the tree file. In all 
analyses, stationarity was reached before one 
million generations, and the first 1,000 trees 
were discarded (i.e., the first million genera- 
tions) from each analysis as the burn-in. To 
obtain the most accurate branch length esti- 
mates possible, the option prset ratepr = vari- 
able was employed per the recommendations 
of Marshall et al. (2006). To evaluate the ef- 
fects that a simpler substitution model could 
have on tree topology, we also conducted 
analyses using the HKY + G model (Hasegawa 
et al., 1985). 

We analyzed protein sequences translated 
from the concatenated 2,310 nt data-set (total 
of 770 a.a., including 143 unambiguously 
aligned Mcox2e a.a.). Finally, we analyzed 
protein sequences translated from the concat- 
enated 1,881 nt data-set (total of 627 a.a.) 
without Mcox2e, again to determine whether 
the inclusion of the extension sequences had 
any effect on the overall topology. For protein 
data-sets, Bayesian searches were run for five 
million generations with eight search chains, 
saving 10,000 trees, and using the Mtrev [prset 
aamodelpr = fixed(mtrev)] substitution model 
(Adachi & Hasegawa, 1996a, b) with the data 
partitioned by gene region. The analyses of 
both protein sequence data-sets reached 
stationarity before 500,000 generations, and 
the first 1,000 trees were discarded as burn- 
in. Thus, for both nucleotide and protein data- 
sets, the Bayesian analyses were run for at 
least 10 times as long as each took to reach 
stationarity. Reliability of the Bayesian topolo- 
gies was evaluated with the posterior prob- 
abilities from the majority-rule consensus 
trees. 

Neighbor-joining analyses of the nucleotide 
and protein data-sets were done using MEGA3 
(Kumar et al., 2004). Reliability of the internal 
nodes of the NJ trees was estimated by 
bootstrapping the data-set with 1,000 replica- 
tions. A branch-and-bound maximum parsi- 
mony search was conducted on the 
concatenated 2,310 nt data-set (which was 
transformed at only the 3" codon positions 
[wherein only transversions were coded]) us- 


308 CHAPMAN ET AL. 


ing PAUP* (version 4.0810: Swofford, 2001). 
A total of 100,000 full heuristic MP bootstrap 
replications was done to estimate the reliabil- 
ity of the internal nodes. 

Differences in topology between constrained 
Bl trees (based on the topologies in Campbell 
et al., 2005, and Zanatta & Murphy, 2006, as 
well as from hypotheses of evolutionary rela- 
tionships deduced from classifications of 
Hannibal, 1912, and Heard & Guckert, 1971) 
and the best BI tree from the unconstrained 
BI analysis of the complete nucleotide data- 
set were tested with the likelihood-based ap- 
proximately unbiased test (AU: Shimodaira, 
2002), the Shimodaira-Hasegawa test (SH: 
Shimodaira & Hasegawa, 1999), the weighted 
Shimodaira-Hasegawa test (WSH: Shimo- 
daira, 2002), the Kishino-Hasegawa test (KH: 
Kishino & Hasegawa, 1989), and the weighted 
Kishino-Hasegawa test (WKH) in CONSEL 
(Shimodaira & Hasegawa, 2001). The program 
Mesquite (version 1.05: Maddison & 
Maddison, 2003) was used to implement maxi- 
mum likelihood estimations of ancestral char- 
acter states on the best BI tree estimated from 
the 2,310 bp data-set under the GTR + G + | 
model. Both the “Markov k-state 1 parameter 
model” (MK1: Lewis, 2001), which assumes 
equally probable forward and backward rates 
of change, and the “Asymmetrical Markov k- 
state 2 parameter model” (AsymmMk: Pagel, 
1997; Mooers & Schluter, 1999), in which “for- 
ward” and “backward” transition rates can be 
different, were investigated. The asymmetry 
likelihood ratio test was used to determine 
whether the AsymmMK model was significantly 
better than the MK1 model (see the Mesquite 
manual). To assess which ancestral charac- 
ter state was best for a given node, state esti- 
mates with a log likelihood 2 or more units 
lower than the best state estimate (decision 
threshold [T] set to T = 2) were rejected 
(Edwards, 1972; Pagel, 1999). Mesquite also 
was used to produce MP-based estimates of 
ancestral character states using the best BI 
tree topology. 


RESULTS AND DISCUSSION 


In the 21 species examined, we obtained 
comparable sequences of the following 
lengths: 672 bp of Fcox1, 279 bp of Fcox2, 
651 bp of Mcox1, and 279 bp of Mcox2h. 
Mcox2e ranged from 543 (Inversidens and 
Amblema) to 561 (Ptychobranchus) bp in 
length (181-187 a.a.). The Mcox2e region 


contains indels within nucleotide positions 7— 
135. The first six nucleotides and those from 
position 136-558 aligned unambiguously and 
were used in analyses containing Mcox2e, 
while those from positions 7-135 were re- 
moved prior to analyses. 

The best tree from the Bayesian analysis of 
the concatenated M and Fcox2-cox1 nucle- 
otide sequences (including Mcox2e) using 
GTR + G + | is shown in Figure 1. Therein, the 
general amblemine tribal relationships indicated 
in Campbell et al. (2005) - e.g., (Quadrulini 
(Pleurobemini (Amblemini, Lampsilini))) — are 
supported but with some notable exceptions 
near the ancestral lineage of the Lampsilini. In 
Figure 1, Popenaias рорей is represented as 
the sister taxon to a clade composed of all of 
the lampsiline species included in this data- 
set. This grouping is supported by a Bayesian 
inference posterior probability (BI PP) of 1.00 
and a MP bootstrap percentage (MP BSP) of 
100. The basal position of P. рорей relative to 
the lampsiline species included herein is sup- 
ported by a BI PP of 1.00 and a MP BSP of 94 
(Fig. 1). Given these high nodal support val- 
ues, this is the first robust demonstration of 
the monophyly of the Lampsilini; both 
Campbell et al. (2005) and Zanatta & Murphy 
(2006) had BI PPs < 0.95 and MP BSPs < 50. 
Furthermore, all of the variations on our Baye- 
sian analyses (including and omitting Mcox2e; 
nt vs. a.a. data) had BI PPs = 0.97 for the 
monophyly of the Lampsilini (Appendix II). The 
inclusion of Amblema and Plectomerus in a 
clade with the traditional lampsiline taxa + 
Popenaias is strongly supported (BI PP = 1.00, 
MP BSP = 96), while their basal placement to 
the traditional lampsiline taxa + Popenaias is 
supported by a BI PP of 1.00 and a MP BSP 
of 100 (Fig. 1). Therefore, the hypothesis that 
Plectomerus and Toxolasma are sister gen- 
era (Campbell et al., 2005: fig. 2) is strongly 
rejected by these results. The paraphyletic 
nature of Amblema + Plectomerus is also sup- 
ported by the present analysis (PP = 0.98, MP 
BSP = 73). The Lampsilis-like species repre- 
sented in this data-set (i.e., L. ovata, L. 
hydiana, L. straminea, Actinonaias ligamentina, 
and Hamiota subangulata) comprise a very 
well-supported (BI PP = 1.00, MP BSP = 100), 
relatively derived clade in the best BI tree 
(Pigs ¥): 

The Bayesian analysis of the five gene re- 
gion data-set using the HKY + G model yielded 
an identical topology. The Bayesian analysis, 
using GTR + | + G, that omitted the Mcox2e 
region was essentially the same, the only dif- 


EVOLUTIONARY RELATIONSHIPS OF POPENAIAS POPEII 


0.99 
Si 


0.99 


1.0 
100 


0.98 
73 


1.0 
96 


0.85 
88 


1.0 
100 


1.0 
100 


Lampsilis straminea 
Lampsilis hydiana 
Actinonaias ligamentina 
Lampsilis ovata 
Hamiota subangulata 


Venustaconcha ellipsiformis 
1.0 
100 


Obovaria olivaria 
Ptychobranchus fasciolaris 
Lemiox rimosus 
Cyrtonaias tampicoensis 
Glebula rotundata 


Obliquaria reflexa 


Toxolasma lividus 


Popenaias popeii 


Plectomerus dombeyanus 


Amblema plicata 


Pleurobema sintoxia 


1.0 
100 


Fusconaia flava 


Pleurobemini 


Quadrula quadrula 


1.0 
100 


Quadrulini 


Quadrula refulgens 


— 0.05 substitutions/site 


309 


Amblemini 


Inversidens japanensis (Gonideini) 


FIG. 1. Bayesian tree, for the 21 amblemine bivalve species, with highest overall posterior probability. 
This tree was generated using the GTR+I+G model on sequences from five mitochondrial gene 
regions, displays maximum likelihood estimated branch lengths and shows nodal support values (PP 
above branch; MP bootstrap percentage below branch [when > 50%]). 


310 


CHAPMAN ET AL. 


TABLE 2. BI constraint analysis results using the five gene region, nucleotide dataset. Only two (in 
parentheses) of the thirty “best tree vs. constraint tree” comparisons were deemed insignificant. 


Tree -In L Difference AU 
Unconstrained 19408.96 (Best) 

Constraint 1 19477.30 68.34 p = 2e-06 
Constraint 2 19431.05 22.10 p=0.014 
Constraint 3 19431.05 22.10 p = 0.014 
Constraint 4 19495.67 86.71 p = 1e-91 
Constraint 5 19460.96 52.00 p=2e-04 
Constraint 6 19488.49 79.54 p = 1e-04 


Constraint 1 = (Plectomerus + Toxolasma) 
Constraint 2 = (Popenaias + Amblema) 


Likelihood-based tests 


KH SH WKH WSH 
р = 1е-04 p=2e-04 p=1e-04 p= 1е-04 
р =0. 019 (p=0.123) p=0.019 p=0.038 
р = 0. 019 (р=0.123) р=0.019 p=0.038 
p=0 p=0 p=0 р = 5e-05 
p=0.001 p=0.001 p=0.001 p=0.001 
р = 4е-05 p=4e-05 p=4e-05 p=4e-05 


( 
| 
((Popenaias, Amblema), ((Plectomerus, Toxolasma), all other lampsilines)) 
( 
( 


Constraint 3 = (Plectomerus, ((Popenaias + Amblema), all lampsilines)) 
Constraint 4 = 

Constraint 5 = (lampsilines + Popenaias + pleurobemines) 

Constraint 6 = (lampsilines + pleurobemines) 


ference being a three-clade polytomy near the 
root that was resolved in Figure 1 (Appendix 
II-A). The analysis of the data-set omitting 
Mcox2e using HKY + G had Amblema and 
Plectomerus exchanging positions, and a 
clade comprised of the two Quadrula species 
was sister to Pleurobema + Fusconaia (Ap- 
pendix II-B). Nevertheless, the evolutionary re- 
lationships indicated by our tree topology in 
Figure 1 are generally similar to those in 
Campbell et al. (2005) and Zanatta & Murphy 
(2006); however, our Figure 1 typically displays 
higher nodal support values. This could be due 
simply to a larger number of informative char- 
acters in our most inclusive nucleotide matrix 
(914 parsimony-informative characters; 
Campbell et al., 2005: 749 parsimony-infor- 
mative characters; Zanatta & Murphy, 2006: 
606 parsimony-informative characters). 

The topology from the BI analysis of the pro- 
tein sequences (Mtrev substitution model) in- 
cluding Mcox2e had only minor differences 
from the tree in Figure 1; Hamiota + Lampsilis 
ovata and (Cyrtonaias + Glebula) + (Ptycho- 
branchus + Lemiox) were paired as sister taxa 
with both pairings exhibiting low nodal support 
(BI PP = 52 and 66, respectively; Appendix II- 
C). The topology from the BI analysis with 
Mcox2e protein sequences deleted had the 
following differences from the topology in Fig- 
ure 1: Actinonaias + Hamiota were paired as 
sister taxa with Lampsilis ovata sister to 
(Hamiota + Actinonaias) + (L. straminea + L. 
hydiana), a clade comprised of the two Qua- 
drula species were sister to Pleurobema + 


Fusconaia and Plectomerus was sister to a 
clade that included all taxa except /nversidens 
(Appendix II-D). 

The neighbor-joining analyses produced es- 
sentially the same tree topologies as Bayesian 
methods, although the bootstrap support for 
some internal nodes was quite low (Appendi- 
ces Ш, IV). The branch-and-bound maximum 
parsimony search on the transformed nucle- 
otide matrix yielded two most parsimonious 
trees (not shown) highly similar to the topol- 
ogy in Figure 1; however, in one, Toxolasma 
lividus and Obliquaria reflexa were paired as 
sister taxa, while the other tree returned 
Obliquaria as sister to ((Lemiox + Ptychobran- 
chus) + (Obovaria + Venustaconcha) + 
(Hamiota + (L. ovata + (Actinonaias + (L. 
straminea+ L. hydiana))))). Of our 11 explicitly 
figured trees, ten displayed Popenaias as the 
sister taxon to a clade containing the traditional 
lampsiline taxa. The exception is a NJ tree 
where Popenaias is sister to Amblema but with 
a NJ BSP of < 50 (Appendix III-D). 

With respect to Amblema, Plectomerus, 
Popenaias, and Toxolasma, results of the con- 
straint analyses (Table 2) strongly suggest that 
the evolutionary relationships implied by our 
best BI tree (Fig. 1) are significantly better than 
those presented in Campbell et al. (2005: figs. 
1,2) and Zanatta & Murphy (2006: fig. 1). Spe- 
cifically, constraint statements #1 (Plectomerus 
+ Toxolasma) and #4 ((Popenaias, Amblema), 
((Plectomerus, Toxolasma), all other lampsili- 
nes)) (Campbell et al., 2005: fig. 2) produced 
trees that were significantly worse than the 


EVOLUTIONARY RELATIONSHIPS OF POPENAIAS POPEII 91 


Location of Marsupium 
L_] Tetragenous 
ШИ Ectobranchous 


Lampsilis hydiana 
Lampsilis straminea 
Actinonaias ligamentina 
Lampsilis ovata 
Hamiota subangulata 
Venustaconcha ellipsiformis 
Obovaria olivaria 
Ptychobranchus fasciolaris 
Lemiox rimosus 
Cyrtonaias tampicoensis 
Glebula rotundata 
Obliquaria reflexa 
Toxolasma lividus 
Popenaias popeii 

|) Plectomerus dombyanus 

|) Amblema plicata 
Pleurobema sintoxia 

U) Fusconaia flava 

[_) Quadrula quadrula 

UL) Quadrula refulgens 


is) Inversidens japanensis 


FIG. 2. ML ancestral character state estimation of the location of the marsu- 
pium, for the 21 amblemine bivalve species, using the Asymmetrical Markov 
k-State 2 parameter model on the tree presented in Figure 1. The asterisk (*) 
denotes the only node that is not significant for the state occupying the ma- 


jority of each pie chart. 


unconstrained tree (Fig. 1) by all five ML test 
statistics (Table 2). Similarly, constraint state- 
ments #2 (Popenaias + Amblema) and #3 
(Plectomerus, ((Popenaias, Amblema), all 
lampsilines)) (Campbell et al., 2005: figs. 1, 2; 
Zanatta & Murphy, 2006: fig. 1) produced trees 
that were deemed significantly worse than the 
unconstrained tree (Fig. 1) by four out of the 
five ML test statistics (Table 2). Constraint 
statements #5 and #6, which evaluated the 
hypotheses that lampsilines have pleuro- 
bemine affinities (e.g., Hannibal, 1912: Heard 
& Guckert, 1971), also produced significantly 
worse topologies than the best unconstrained 
tree (Fig. 1). 


Figure 1 is demonstrably the best current 
estimate of phylogeny for the present taxa/data 
combination. Using this tree, ML ancestral 
character state estimation procedures gener- 
ated inferences regarding evolutionary transi- 
tions in the location of the marsupium for the 
included species (Fig. 2). For this character 
state optimization, the AsymmMK model was 
significantly better than the MK1 model and 
the former was used to optimize the location 
of the marsupium onto the tree in Figure 1. 
The ancestral character state estimations for 
all of the internal nodes of the tree were sig- 
nificant for the majority state present in each 
pie chart except for the node leading to 


312 CHAPMAN ET AL. 


Popenaias (denoted by an asterisk). The MP 
estimate (not shown) was essentially identical 
to that from ML. Figure 2 indicates that the ecto- 
branchy present in the “traditional” lampsiline 
taxa + Popenaias was derived a single time 
from the tetragenous brooding condition found 
in an Amblema/Plectomerus-like ancestor 
rather than from the plesiomorphic retention 
of pleurobemine ectobranchy. It is readily ap- 
parent that the evolution of ectobranchy was a 
necessary antecedent to the attainment of the 
specialized “heterogenae” brooding condition 
(i.e., posteriorly located marsupia in only the 
outer demibranchs) now found in the more 
derived lampsiline taxa. 

The presence of a principally ectobranchous 
brooding location in Popenaias, the topologies 
presented in Figure 1 and Appendices II-IV and 
the constraint analysis results (Table 2) sug- 
gest that Popenaias has greater propinquity to 
the “traditional” lampsiline genera than to 
Amblema or Plectomerus. The phylogenetic 
placement of Popenaias popeil in Figure 1 and 
the occasional use of its inner demibranchs for 
brooding suggest that this species likely pos- 
sesses some characteristics of the lampsiline 
ancestral lineage. Thus, further studies of 
Popenaias, Amblema, and Plectomerus mor- 
phology, reproduction, ecology and phylogeo- 
graphy combined with that of other pertinent 
taxa — e.g., “Fusconaia” ebena (Lea 1831), 
“Obovaria” rotulata (Wright 1899), Actinonaias 
sapotalensis (Lea 1841), Friersonia iridella 
(Pilsbry & Frierson 1908), and Nephronaias, 
5.5., spp. (Ortmann, 1912; Simpson, 1914; 
Fuller, 1975; Campbell et al., 2005) — are es- 
sential to form a better understanding of the 
evolutionary antecedents to the extraordinary 
morphological radiation that occurred during 
lampsiline phylogenesis. 

As mentioned in the Introduction, a narrow 
use of only the morphological “lampsiline di- 
agnostic” character states would likely result 
in the exclusion of some species typically per- 
ceived to be “lampsiline.” However, recent phy- 
logenetic analyses utilizing molecular data 
have supported the monophyly of the traditional 
lampsiline species (e.g., Campbell et al., 2005; 
Zanatta & Murphy, 2006; trees presented 
herein), which strongly attests to the reality of 
the group. Given this situation and the lack of 
obvious patristic distance-based “gaps” near 
the base of the lampsiline/Popenaias/Plecto- 
merus/Amblema clade in Figure 1, delimiting 
“What is a lampsiline?” herein would consti- 
tute a relatively subjective decision. Instead, 
we suggest the use of the tribal designation 


Amblemini for the lampsiline/Popenaias/Plecto- 
merus/Amblema clade in Figure 1 because the 
usage of Amblemini Rafinesque, 1820, in a 
tribal context has priority over Lampsilini 
Ihering, 1901. This strategy eliminates the pro- 
duction of a paraphyletic Amblemini group, 
which would likely contain at least Plectomerus 
and Amblema, if a restricted lampsiline clade 
within the more inclusive group was designated 
as the Lampsilini. Furthermore, this implemen- 
tation does not preclude a future recognition 
of a distinct lampsiline clade at the subtribal 
rank (i.e., Lampsilina). 

Due to issues involving limited taxon sam- 
pling, it is readily apparent that the evolution- 
ary relationships presented in this study are 
not conclusive regarding amblemine evolution- 
ary relationships. However, the general agree- 
ment of our amblemine relationships, based 
on total molecular evidence analyses of se- 
quences from two independent mtDNA ge- 
nomes with those depicted in more 
taxonomically inclusive, F mtDNA genome- 
only studies (e.g., Campbell et al., 2005; 
Zanatta & Murphy, 2006) suggests that the 
broad outline of amblemine phylogeny has 
emerged. Furthermore, it should be noted that 
most of the parsimony-informative characters 
in the analyses presented herein came from 
the M mtDNA sequences, further illuminating 
the potential importance of M genomes in 
freshwater mussel phylogenetics (Hoeh et al., 
2002; Walker et al., 2006). Subsequent stud- 
ies of unionoidean bivalve evolution will ben- 
efit greatly from combined analyses of F 
mtDNA-, М mtDNA- and nucleus-encoded 
DNA sequences as well as the inclusion of 
morphological and behavioral characters. 


ACKNOWLEDGMENTS 


We thank $. A. Ahlstedt, С. Barnhart, А. E. 
Bogan, К. $. Butler, А. D. Christian, J. Е. Har- 
ris, W. H. Heard, R. G. Howells, D. Hubbs, J. 
W. Jones, P. Morrison, W. R. Posey, B. D. 
Sietman, R. J. Trdan and G. Zimmerman for 
providing specimens and D. Senyo for assis- 
tance in the lab. We also thank G. M. Davis, 
W. Н. Heard, D. L. Graf and two anonymous 
reviewers for suggestions that led to a signifi- 
cantly improved manuscript. This work was 
supported by grants from the National Science 
Foundation (DEB-0237175 to W.R.H.), the 
Division of Federal Aid of the US Fish & Wild- 
life Service (Region 2) and the New Mexico 
Department of Game and Fish. 


EVOLUTIONARY RELATIONSHIPS OF POPENAIAS РОРЕП 313 


LITERATURE CITED 


ADACHI, J. & M. HASEGAWA, 1996a, Model of 
amino acid substitution in proteins encoded by 
mitochondrial DNA. Journal of Molecular Evo- 
lution, 42: 459-468. 

ADACHI, J. & М. HASEGAWA, 1996b, MOLPHY 
version 2.3: programs for molecular phylo- 
genetics based on maximum likelihood. /nsti- 
tute of Statistical Mathematics Computer 
Science Monographs, 28: 1-150. 

BRETON, S., H. D. BEAUPRE, О. T. STEWART, 
W.H. HOEH & P. U. BLIER, 2007, The unusual 
system of doubly uniparental inheritance of 
mtDNA: isn’t one enough? Trends in Genet- 
ics, 23: 465-474. 

BURCH, J., 1975, Freshwater unionacean clams 
(Mollusca: Pelecypoda) of North America. 
Hamburg, Malacological Publications. 204 pp. 

CAMPBELL, D.-C., J.M. SERB, J. Е. BUHAY, К. 
J. ROE, R. L. MINTON & С. LYDEARD, 2005, 
Phylogeny of North American amblemines 
(Bivalvia, Unionoida): prodigious polyphyly 
proves pervasive across genera. Invertebrate 
Biology, 124: 131-164. 

CHAKRABARTI R., J. M. WALKER, Е. С. 
CHAPMAN, S. SHEPARDSON, R. J. TRDAN, 
J.P. CUROLE, G. T. WATTERS, О.Т. STEWART, 
S. VIJATRAGHAVAN & W. R. HOEH, 2007, 
Reproductive function for a c-terminus ex- 
tended, male-transmitted cytochrome c oxi- 
dase subunit Il protein expressed in both 
spermatozoa and eggs. FEBS Letters, 581: 
5213-5219. 

CHAKRABARTI, R., J. M. WALKER, D. T. STEW- 
ART, R. J. TRDAN, $. VIJATRAGHAVAN, J. 
CUROLE & W. В. НОЕН, 2006, Presence of a 
unique male-specific extension of C-terminus 
to the cytochrome с oxidase subunit II protein 
coded by the male-transmitted mitochondrial 
genome of Venustaconcha ellipsiformis (Bi- 
valvia: Unionoidea). FEBS Letters, 580: 862- 
866. 

CHAMBERLAIN, T., 1934, The glochidial con- 
glutinates of the Arkansas fanshell, Cyprogenia 
aberti (Conrad). Biological Bulletin, 66: 55-61. 

COCKERELL, Т. D. A., 1902, Unio popei, Lea, 
in New Mexico. The Nautilus, 16: 69-70. 

CUROLE, J. P. & T. D. KOCHER, 2002, Ancient 
sex-specific extension of the cytochrome c 
oxidase II gene in bivalves and the fidelity of 
doubly-uniparental inheritance. Molecular Bi- 
ology and Evolution, 19: 1323-1328. 

CUROLE, J. P. 4 Т. О. KOCHER, 2005, Evolu- 
tion of a unique mitotype-specific protein-cod- 
ing extension of the cytochrome с oxidase ll 
gene in freshwater mussels (Bivalvia: 
Unionoida). Journal of Molecular Evolution, 61: 
381-389. 

DAVIS, С. M. 8 S. L. H. FULLER, 1981, Genetic 
relationships among recent Unionacea (Bi- 
valvia) of North America. Malacologia, 20: 217- 
253; 

EDWARDS, A. W. F., 1972, Likelihood. Cam- 
bridge, Cambridge University Press. 252 pp. 
FOLMER, O., M. BLACK, W. R. HOEH, R. LURZ 
& К. С. VRIJENHOEK, 1994, DNA primers for 
amplification of mitochondrial cytochrome c 


oxidase subunit | from diverse metazoan in- 
vertebrates. Molecular Marine Biology and Bio- 
technology, 3: 294-299. 

FULLER, S. L. H., 1975, The systematic posi- 
tion of Cyrtonaias (Bivalvia, Unionidae). Mala- 
cological Review, 8: 81-89. 

FRIERSON, L. S., 1927, A classified and anno- 
tated check list of the North American naiades. 
Waco, Texas, Baylor University Press. 111 pp. 

GRAF, D. L. & D. O FOIGHIL, 2000, The evolu- 
tion of brooding characters among the fresh- 
water pearly mussels (Bivalvia: Unionoidea) of 
North America. Journal of Molluscan Studies, 
66: 157-170. 

HAAG, W. R., R. S. BUTLER & P. D. HARTFIELD, 
1995, An extraordinary reproductive strategy 
in freshwater bivalves: prey mimicry to facili- 
tate larval dispersal. Freshwater Biology, 34: 
471-476. 

HANNIBAL, H., 1912, A synopsis of the recent 
and Tertiary freshwater Mollusca of the Cali- 
fornian Province, based upon an ontogenetic 
classification. Proceedings of the Malacologi- 
cal Society of London, 10: 112-211. 

HASEGAWA, M., К. KISHINO & T. YANO, 1985, 
Dating the human-ape splitting by a molecular 
clock of mitochondrial DNA. Journal of Molecu- 
lar Evolution, 22: 160-174. 

HEARD, W. H., 1974, Anatomical systematics of 
freshwater mussels. Malacological Review, 7: 
41-42. 

HEARD, W. H. & R. H. GUCKERT, 1971, A re- 
evaluation of the recent Unionacea 
(Pelecypoda) of North America. Malacologia, 
10: 333-335. 

HINCKLEY,A.A., 1907, Shells collected in north- 
eastern Mexico. The Nautilus, 21: 68-72, 76- 
80. 

НОЕН, W.R., О. T. STEWART & $. I. GUTTMAN, 
2002, High fidelity of mitochondrial genome 
transmission under the doubly uniparental 
mode of inheritance in freshwater mussels 
(Bivalvia: Unionoidea). Evolution, 56: 2252- 
2261. 

НОЕН, W.R., О. T. STEWART, В. SUTHERLAND 
& E. ZOUROS, 1996, Multiple origins of gen- 
der associated mitochondrial DNA lineages in 
bivalves (Mollusca: Bivalvia). Evolution, 50: 
2276-2286. 

HUELSENBECK, J. P. & F. RONQUIST, 2005, 
Bayesian analysis of molecular evolution us- 
ing MrBayes. Pp. 183-222, in: R. NIELSEN, ed., 
Statistical methods in molecular evolution. New 
York, Springer. 504 pp. 

IHERING, H. VON, 1901, The Unionidae of North 
America. The Nautilus, 15: 37-39, 50-53. 

JOHNSON, R. 1., 1972, The Unionidae (Mollusca: 
Bivalvia) of peninsular Florida. Bulletin of the 
Florida State Museum of Natural History, 16: 
181-249. 

JOHNSON, К. I., 1999, The Unionidae of the Rio 
Grande (Rio Bravo del Norte) system of Texas 
and Mexico. Occasion Papers on Mollusks, 77: 
1-65. 

JONES, J. W., R. J. NEVES, S.A. AHLSTEDT & 
R.A. MAIR, 2004, Life history and propagation 
of the endangered dromedary pearlymussel 
(Dromus dromas) (Bivalvia: Unionidae). Jour- 


314 CHAPMAN ETAL. 


nal of the North American Benthological Soci- 
ety, 23: 515-525. 

KISHINO, H. & M. HASEGAWA, 1989, Evalua- 
tion of the maximum likelihood estimate of the 
evolutionary tree topologies from DNA se- 
quence data, and the branching order of the 
Hominoidea. Journal of Molecular Evolution, 
29: 170-179. 

KREBS, R. A., 2004, Combining paternally and 
maternally inherited mitochondrial DNA for 
analysis of population structure in mussels. 
Molecular Ecology, 13: 1701-1705. 

KUMAR, S., K. TAMURA & M. NEI, 2004, 
МЕСАЗ: integrated software for molecular evo- 
lutionary genetics analysis and sequence align- 
ment. Briefings in Bioinformatics, 5: 150-163. 

LEWIS, P. O., 2001, A likelihood approach to 
estimating phylogeny from discrete morpho- 
logical character data. Systematic Biology, 50: 
913-925. 

LIU, H.-P., 4. В. MITTON & S.-K. WU, 1996, Pa- 
ternal mitochondrial DNA differentiation far 
exceeds maternal mitochondrial DNA and 
allozyme differentiation in the freshwater mus- 
sel, Anodonta grandis grandis. Evolution, 50: 
952-957. 

LYDEARD, C., M. MULVEY & G. M. DAVIS, 1996, 
Molecular systematics and evolution of repro- 
ductive traits of North American freshwater 
unionacean mussels (Mollusca: Bivalvia) as 
inferred from 16S rRNA gene sequences. 
Philosophical Transactions of the Royal Soci- 
ety London, (B), 351: 1593-1603. 

MADDISON, W. P. & D. R. MADDISON, 2000, 
MacClade 4: analysis of phylogeny and char- 
acter evolution. Sunderland, Sinauer Associ- 
ates, Inc. CD-ROM. 

MADDISON, W. P. & D. R. MADDISON, 2003, 
Mesquite: a modular system for evolutionary 
analysis, version 1.05: http://mesquiteproject.org 

MARSHALL, D. C., С. SIMON & Т. В. BUCKLEY, 
2006, Accurate branch length estimation in 
partitioned Bayesian analyses requires accom- 
modation of among-partition rate variation and 
attention to branch length priors. Systematic 
Biology, 55: 992-1003. 

MIZI, A., Е. ZOUROS, М. MOSCHONAS & С. С. 
RODAKIS, 2005, The complete maternal and 
paternal mitochondrial genomes of the Medi- 
terranean mussel Mytilus galloprovincialis: 
implications for the doubly uniparental inherit- 
ance mode of mtDNA. Molecular Biology and 
Evolution, 22: 952-967. 

MOOERS, A: -O. -& D: :SCHEUTER, 1999, "Re- 
constructing ancestor states with maximum 
likelihood: support for one- and two-rate mod- 
els. Systematic Biology, 48: 623-633. 

NEI, M. & $. KUMAR, 2000, Molecular evolution 
and phylogenetics. Oxford, Oxford University 
Press. 333 pp. 

OBATA, M., N. SANO, K. KAWAMURA & A. 
KOMARU, 2007, Inheritance of two M type 
mitochondrial DNA from sperm and unfertilized 
eggs to offspring in Mytilus galloprovincialis. 
Development, Growth and Differentiation, 49: 
335-344. 

ORTMANN, A. E., 1910, A new system of the 
Unionidae. The Nautilus, 23: 114-120. 


ORTMANN, A. E., 1912, Notes upon the fami- 
lies and genera of the najades. Annals of the 
Carnegie Museum, 8: 222-365. 

PAGEL, M., 1997, Inferring evolutionary pro- 
cesses from phylogenies. Zoologica Scripta, 
26: 331-348. 

PAGEL, M., 1999, The maximum likelihood ap- 
proach to reconstructing ancestral character 
states on phylogenies. Systematic Biology, 48: 
612-622. 

PILSBRY, H. A., 1909a, Unionidae from the 
Panuco River system, Mexico. Proceedings of 
the Academy of Natural Sciences of Philadel- 
phia, 61: 532-539. 

PILSBRY, H.A., 1909b, Mollusks from northeast- 
ern Mexico. The Nautilus, 23: 45-49. 

POSADA, D. & K. A. CRANDALL, 1998, 
Modeltest: testing the model of DNA substitu- 
tion. Bioinformatics, 14: 817-818. 

RAWSON, Р. D. & Т. J. HILBISH, 1995, Evolu- 
tionary relationships among the male and fe- 
male mitochondrial DNA lineages in the Mytilus 
edulis species complex. Molecular Biology and 
Evolution, 12: 893-901. 

RODRIGUEZ, F., J. L. OLIVER, A. MARIN & J. 
R. MEDINA, 1990, The general stochastic 
model of nucleotide substitution. Journal of 
Theoretical Biology, 142: 485-501. 

ROE, K. J. & P. HARTFIELD, 2005, Hamiota, a 
new genus of freshwater mussel (Bivalvia: 
Unionidae) from the southeastern United 
States. The Nautilus, 119: 1-10. 

RONQUIST, Е. & J. P. HUELSENBECK, 2003, 
MrBayes 3: Bayesian phylogenetic inference 
under mixed models. Bioinformatics, 19: 1572- 
1574. 

SHIMODAIRA, H., 2002, An approximately un- 
biased test of phylogenetic tree selection. Sys- 
tematic Biology, 51: 492-508. 

SHIMODAIRA, H. & M. HASEGAWA, 1999, Mul- 
tiple comparisons of log-likelihoods with appli- 
cations to phylogenetic inference. Molecular 
Biology and Evolution, 16: 1114-1116. 

SHIMODAIRA, H. & M. HASEGAWA, 2001, 
CONSEL: for assessing the confidence of phy- 
logenetic tree selection. Bioinformatics, 17: 
1246-1247. 

SIMPSON, С. Т., 1900, Synopsis of the naiades, 
or pearly fresh-water mussels. Proceedings of 
the United States National Museum, 32: 501- 
1044. 

SIMPSON, С. T., 1914, A descriptive catalogue 
of the naiades or pearly fresh-water mussels. 
Detroit, Michigan, Bryant Walker. 1540 pp. 

SKIBINSKI, D. O. F., С. GALLAGHER & С. M. 
BEYNON, 1994, Sex-limited mitochondrial 
DNA transmission in the marine mussel Mytilus 
edulis. Genetics, 138: 801-809. 

SMITH, D. С., В. К. LANG & M. Е. GORDON, 
2003, Gametogenetic cycle, reproductive 
anatomy, and larval morphology of Popenaias 
popeii (Unionoida) from the Black River, New 
Mexico. The Southwestern Naturalist, 48: 333- 
340. 

STEWART, D. T., C. SAAVEDRA, R. R. 
STANWOOD, A. BALL & E. ZOUROS, 1995, 
Male and female mitochondrial DNA lineages 
in the Blue mussel (Mytilus edulis) species 


EVOLUTIONARY RELATIONSHIPS OF POPENAIAS POPEII 319 


group. Molecular Biology and Evolution, 12: 
735-747. 

SUTHERLAND: .B;, D, SLEWART ER. 
KENCHINGTON & Е. ZOUROS, 1998, The fate 
of paternal mitochondrial DNA in developing 
female mussels Mytilus edulis: implications for 
the mechanism of doubly uniparental inherit- 
ance of mitochondrial DNA. Genetics, 148: 
341-347. 

SWOFFORD, D. L., 2002, PAUP*. Phylogenetic 
Analysis Using Parsimony (and other methods) 
4.0 Beta. Sunderland, Sinauer Associates, Inc. 
CD-ROM. 

TURGEON, D. D., J. R. QUINN, A. E. BOGAN, E. 
V. COAN, Е. С. HOCHBERG, W. С. LYONS, P. 
М. MIKKELSEN, К. J. NEVES, С.Е. Е. ROPER, 
G. ROSENBERG, B. ROTH, A. SCHELTEMA, 
F. G. THOMPSON, M. VECCHINOE & J. D. WIL- 
LIAMS, 1998, Common and scientific names of 
aquatic invertebrates from the United States and 
Canada: mollusks. (second edition). American 
Fisheries Society Special Publication, 26: 1-526. 

WALKER, УМ, JP. CUROLE „ВЕ WADE. Е. 
С. CHAPMAN, А. Е. BOGAN, С. Т. WATTERS 
& W. К. НОЕН, 2006, Taxonomic distribution 


and phylogenetic utility of gender-associated 
mitochondrial genomes in the Unionoida 
(Bivalvia). Malacologia, 48: 265-282. 

WALKER, J. M., A. E. BOGAN, Е. A. BONFIGLIO, 
D. C. CAMPBELL, A. D. CHRISTIAN, J. P. 
CUROLE, J. L. HARRIS, R. J. WOJTECKI & 
W. R. HOEH, 2007, Primers for amplifying the 
hypervariable, male-transmitted COII-COl 
junction region in amblemine freshwater mus- 
sels (Bivalvia: Unionoidea: Ambleminae). Mo- 
lecular Ecology Notes, 7: 489-491. 

WATTERS, G. T., 2001, The evolution of the 
Unionacea in North America, and its implica- 
tions for the worldwide fauna. Pp. 281-307, in: 
G. BAUER & K. WACHTLER, eds., Ecology and 
evolution of the freshwater mussels 
(Unionoida). Berlin, Springer-Verlag. 394 pp. 

ZANATTA, О. Т. & К. W. MURPHY, 2006, Evolu- 
tion of active host-attraction strategies in the 
freshwater mussel tribe Lampsilini (Bivalvia: 
Unionidae). Molecular Phylogenetics and Evo- 
lution, 41: 195-208. 


Revised ms. accepted 12 March 2008 


APPENDIX | 


Collection locality information for the amblemine bivalve specimens sequenced herein. (Spe- 


cies authors and dates given in Table 1.) 


Quadrula quadrula — East Fork of the White 
River, downstream of US 50/SR 37 bridge, 
Lawrence County, Indiana. 

Quadrula refulgens — Amite River, near Port 
Vincent, East Baton Rouge Parish, Louisiana. 

Fusconaia flava — Ohio River @ River Mile 625, 
near Fishtown, Harrison County, Indiana. 

Pleurobema sintoxia — East Fork of the White 
River, downstream of US 50/SR 37 bridge, 
Lawrence County, Indiana. 

Actinonaias ligamentina — French Creek @ 
Gravel Run Road crossing, Venango, Craw- 
ford County, Pennsylvania. 

Amblema plicata — Black River @ Aitken Road 
crossing, Sanilac County, Michigan. 

Cyrtonaias tampicoensis — Guadalupe River, 
upstream of Wood Lake and upstream of the 
access road from the town of Cost, Gonzales 
County, Texas. 

Glebula rotundata — Bayou Carron at State 
Highway 10 crossing, near town of Washing- 
ton, St. Landry Parish, Louisiana. 

Hamiota subangulata — Tributary of Lower Flint 
River (Spring Creek) @ US 84 crossing, 
Decatur County, Georgia. 

Lampsilis hydiana — Cossatot River @ State High- 
way 24 crossing, Sevier County, Arkansas. 


Lampsilis ovata — Ohio River @ River Mile 726, 
Hancock County, Kentucky. 

Lampsilis straminea — Brush Creek (Tombigbee 
River tributary) @ State Highway 14 cross- 
ing, Greene County, Alabama. 

Lemiox rimosus — Duck River @ River Mile 
179.1, Milltown, Marshall County, Tennessee. 

Obliquaria reflexa — Duck River @ River Mile 
179.1, Milltown, Marshall County, Tennessee. 

Obovaria olivaria — St. Croix River at Interstate 
Park, near Taylors Falls, Chisago County, Min- 
nesota/Polk County, Wisconsin. 

Plectomerus dombeyanus — Ouachita К. @ 
Highway 79B crossing, Camden, Ouachita 
County, Arkansas. 

Popenaias рорей — Black River (Pecos River 
drainage), Eddy County, New Mexico. 

Ptychobranchus fasciolaris — French Creek @ 
Gravel Run Road bridge, Venango, Crawford 
County, Pennsylvania. 

Toxolasma lividus — Clinton River, outflow of 
Dawsons Mill Pond, Beaudette Park, up- 
stream of Orchard Lake Road crossing, 
Pontiac, Oakland County, Michigan. 

Venustaconcha ellipsiformis — Sugar River, 
upstream of state Highway M-30 crossing, 
Gladwin County, Michigan. 


3116 


CHAPMAN ET AL. 


APPENDIX II 


Consensus trees of 21 amblemine bivalve taxa from Bayesian analyses of (A) the nucleotide 
dataset excluding the Mcox2 extension using the GTR+G+I model; (В) the nucleotide dataset 
excluding the Mcox2 extension using the HKY+G model; (C) the amino acid dataset including 
the extension using the Mtrev model; (D) the amino acid dataset excluding the extension using 
the Mtrev model. Numbers above the branches are posterior probabilities x 100. 


A 


100 
100 
88 
100 
100 
100 
98 
57 100 
98 100 
98 
100 
65 
100 
100 
100 
100 
100 
100 
52 
100 
100 
100 
100 
99 66 
100 
99 
100 
62 
100 
62 
100 
100 


Lampsilis straminea B 
Lampsilis hydiana 
Actinonaias ligamentina 
Lampsilis ovata 

Hamiota subangulata 
Venustaconcha ellipsiformis 
Obovaria olivaria 
Ptychobranchus fasciolaris 


Lemiox rimosus 
Cyrtonaias tampicoensis 
Glebula rotundata 
Obliquaria reflexa 
Toxolasma lividus 
Popenaias popeii 
Plectomerus dombeyanus 
Amblema plicata 
Pleurobema sintoxia 
Fusconaia flava 
Quadrula quadrula 
Quadrula refulgens 


Inversidens japanensis 


Lampsilis straminea D 
Lampsilis hydiana 
Actinonaias ligamentina 
Lampsilis ovata 

Hamiota subangulata 
Venustaconcha ellipsiformis 
Obovaria olivaria 
Cyrtonaias tampicoensis 
Glebula rotundata 
Ptychobranchus fasciolaris 
Lemiox rimosus 

Obliquaria reflexa 
Toxolasma lividus 
Popenaias popeii 
Plectomerus dombeyanus 
Amblema plicata 
Pleurobema sintoxia 
Fusconaia flava 

Quadrula quadrula 
Quadrula refulgens 


Inversidens japanensis 


99 


100 
100 
91 
100 
100 
100 
99 
a 100 
22 100 
70 
100 
97 
100 
67 
10 
100 
93 
99 80 
100 
70 100 
98 100 
100 100 
97 
97 
99 
60 
100 
100 

100 


Lampsilis straminea 
Lampsilis hydiana 
Actinonaias ligamentina 
Lampsilis ovata 

Hamiota subangulata 
Venustaconcha ellipsiformis 
Obovaria olivaria 
Ptychobranchus fasciolaris 
Lemiox rimosus 
Cyrtonaias tampicoensis 
Glebula rotundata 
Obliquaria reflexa 
Toxolasma lividus 
Popenaias popeii 
Amblema plicata 
Plectomerus dombeyanus 
Pleurobema sintoxia 
Fusconaia flava 

Quadrula quadrula 
Quadrula refulgens 


Inversidens japanensis 


Lampsilis straminea 
Lampsilis hydiana 
Actinonaias ligamentina 
Hamiota subangulata 
Lampsilis ovata 
Venustaconcha ellipsiformis 
Obovaria olivaria 
Ptychobranchus fasciolaris 
Lemiox rimosus 
Cyrtonaias tampicoensis 
Glebula rotundata 
Obliquaria reflexa 
Toxolasma lividus 
Popenaias popeii 
Amblema plicata 
Pleurobema sintoxia 
Fusconaia flava 

Quadrula quadrula 
Quadrula refulgens 
Plectomerus dombeyanus 


Inversidens japanensis 


EVOLUTIONARY RELATIONSHIPS OF POPENAIAS POPEII 317 


APPENDIX Ill 


Neighbor-joining phylogenetic trees of 21 amblemine bivalve taxa. Bootstrap values are based 
on 1,000 replications, and only those greater than 50% are shown. /nversidens japanensis was 
used as outgroup. (A): Analysis based on concatenated partial amino acid sequences of Mcox7, 
Mcox2, Fcox1, Fcox2 and Mcox2e. Dayhoff's distance and complete-deletion option were used 
(628 shared sites); (B): Analysis based on concatenated partial nucleotide sequences of Mcox7, 
Mcox2, Fcox1, Fcox2 and Mcox2e. Tamura-Nei distance and complete-deletion option were 
used (1895 shared sites); (C): Analysis based on concatenated partial amino acid sequences of 
Mcox1, Mcox2, Fcox1, Fcox2 (without Mcox2e). Dayhoff's distance and complete-deletion op- 
tion were used (485 shared sites); (D): Analysis based on concatenated partial nucleotide se- 
quences of Mcox1, Mcox2, Fcox1, Fcox2 (without Mcox2e). Tamura-Nei distance and com- 
plete-deletion option were used (1,466 shared sites). 


A ve h B 100 —Lampsilis straminea 
100rLampsilis straminea 100 ne 
100] !Lampsilis hydiana 90 ee : 
22 Actinonaias ligamentina egestas CASE 
100 1: e me Lampsilis ovata 
FRS 100 Hamiota subangulata 
100 Hamiota subangulata E —_ р 
100 Venustaconcha ellipsiformis 70 100 Venustaconcha ellipsiformis 
Obovaria olivaria BENIN TA 
100 Eyrionalastampieoensis 100 Ptychobranchus fasciolaris 
Glebula rotundata Estos fie am 
= 100 Ptychobranchus fasciolaris 891.99 Cyrtonaias tampicoensis 
nuire Glebula rotundata 
96 Obliquaria reflexa 95 Obliquaria reflexa 
60 oc dt 94 53 Toxolasma lividus q 
50 Popenaias popeii Popenaias рорей 
Amblema plicata Plectomerus dombeyanus 
Plectomerus dombeyanus 50 Amblema plicata 
100 Pleurobema sintoxia 100 Pleurobema sintoxia 
97 Fusconaia flava Fusconaia flava 
100 Quadrula quadrula 62 100 Quadrula quadrula 
| Quadrula refulgens Quadrula refulgens 
Inversidens japanensis Inversidens japanensis 
0.02 0.05 
E 93 y Lampsilis straminea D 100 —Lampsilis straminea 
a 25 Lampsilis hydiana 100 Lampsilis hydiana 
a Astingnalas ligamentina 76 Actinonaias ligamentina 
Lampsilis ovata 66 Lampsilis ovata 
99 Hamiota subangulata 92 Hamiota subangulata 
100 Venustaconcha ellipsiformis 100 Venustaconcha ellipsiformis 
58 Obovaria olivaria Obovaria olivaria 
94 Cyrtonaias tampicoensis 100 Ptychobranchus fasciolaris 
70 Glebula rotundata Lemiox rimosus 
57 99 Ptychobranchus fasciolaris 58 || 94 Cyrtonaias tampicoensis 
da Lemiox rimosus 70 Glebula rotundata 
Obliquaria reflexa Obliquaria reflexa 
69 Toxolasma lividus 53 Toxolasma lividus 
Popenaias popeii 79 Popenaias popeii 
Amblema plicata Amblema plicata 
Plectomerus dombeyanus Plectomerus dombeyanus 
99 Pleurobema sintoxia 100 Pleurobema sintoxia 
79 Fusconaia flava Fusconaia flava 
100 Quadrula quadrula 54 100 Quadrula quadrula 
| Quadrula refulgens | Quadrula refulgens 
Inversidens japanensis Inversidens japanensis 
0.01 0.02 


318 CHAPMAN ET AL. 
APPENDIX IV 


Neighbor-joining phylogenetic trees of 21 amblemine bivalve taxa. Bootstrap values are based 
on 1,000 replications, and only those greater than 50% are shown. Inversidens japanensis was 
used as outgroup. (A): Analysis based on partial amino acid Mcox2e sequences. Dayhoff's 
distance and complete-deletion option were used (143 shared sites); (B): Analysis based on 
Mcox2e nucleotide sequences. Tamura-Nei distance and complete-deletion option were used 
(429 shared sites). Dayhoff’s distance, computed as Poisson-corrected gamma distance with a 
= 2.25 (Nei & Kumar, 2000), was used to take into account parallel and backward substitutions. 


A 98 y Lampsilis straminea B 100r Lampsilis straminea 
1001 I Lampsilis hydiana 100 Lampsilis hydiana 
52 Actinonaias ligamentina 58 Actinonaias ligamentina 
100 Lampsilis ovata _100 Lampsilis ovata 
100 Hamiota subangulata 100 Hamiota subangulata 
100 Venustaconcha ellipsiformis 100 Venustaconcha ellipsiformis 
Obovaria olivaria Obovaria olivaria 
5961 Toxolasma lividus Obliquaria reflexa 
94 Cyrtonaias tampicoensis 54 Toxolasma lividus 
Glebula rotundata 94 Cyrtonaias tampicoensis 
Obliquaria reflexa Glebula rotundata 
85 95 Ptychobranchus fasciolaris 2 98 Ptychobranchus fasciolaris 
Lemiox rimosus 69 Lemiox rimosus 

Popenaias popeii Popenaias popeii 

Amblema plicata | Plectomerus dombeyanus 
100r Pleurobema sintoxia 50 Amblema plicata 
75 Fusconaia flava 100 Pleurobema sintoxia 
100 Quadrula quadrula Fusconaia flava 

Quadrula refulgens 100 Quadrula quadrula 
| Plectomerus dombeyanus | Quadrula refulgens 
Inversidens japanensis Inversidens japanensis 
rs | (==) 


0.1 0.05 


RESEARCH NOTES 


MALACOLOGIA, 2008, 50(1-2): 321-330 


CHILINA IGUAZUENSIS (GASTROPODA: CHILINIDAE), 
NEW SPECIES FROM IGUAZU NATIONAL PARK, ARGENTINA 


Diego E. Gutiérrez Gregoric* & Alejandra Rumi 


CONICET, Divisiön Zoologia Invertebrados, Museo de La Plata, Facultad de Ciencias 
Naturales y Museo, Universidad Nacional de La Plata, La Plata, Buenos Aires, Argentina 


ABSTRACT 


The Chilinidae comprises 16 species currently cited for Argentina, mostly distributed in 
Patagonia. All original descriptions of these species have been based on shell characters, 
and their internal anatomy is poorly known. Here a new species, Chilina iguazuensis, is 
described, including shell, radula, and reproductive and nervous systems. This species, 
found in Iguazu National Park, Misiones Province, Argentina, in the Upper Iguazu River 
rapids, has the following distinctive characteristics: aperture length equal to last whorl 
length; central radular tooth asymmetric and bicuspid, with both cusps serrated; and pre- 
puce length 60% of penis sheath length. Chilina iguazuensis is endemic in Iguazú Na- 
tional Park, along with Chilina megastoma, which inhabits waterfalls in the same river. 

Key words: Chilina iguazuensis, n. sp., Chilinidae, anatomy, Argentina. 


INTRODUCTION 


The family Chilinidae Dall, 1870 (Gastropoda, 
Pulmonata, Basommatophora), is exclusive to 
South America, occurring from the Tropic of 
Capricorn to Cape Horn and the Falkland Is- 
lands. The family comprises a single genus, 
Chilina Gray, 1828 (type species Auricula 
(Chilina) fluctuosa Gray, 1828, Chile), with 
about 32 species, 16 of which have been re- 
corded in Argentina (Castellanos & Miquel, 
1980, 1991; Castellanos & Gaillard, 1981; 
Castellanos & Landoni, 1995). Original species 
descriptions are restricted to shell characters 
(Hidalgo, 1880; Marshall, 1924, 1933; Hylton 
Scott, 1958, among others), and very few sub- 
sequent works have dealt with internal anatomy 
(Haeckel, 1911; Duncan, 1960a, b, 1975; Harry, 
1964; Brace, 1983; ltuarte, 1997). Miquel (1984, 
1987) analyzed the penial complex of eight 
species from Argentina, Brazil, Chile, and Uru- 
guay, and concluded that no microanatomical 
differences could be found at this level. Based 
on characteristics of Chilean species, 
Valdovinos & Stuardo (1995) rearranged the 
family systematics and proposed that the ner- 
vous system had the only reliable characters 
enabling definitions of supraspecific taxa. 

From an evolutionary perspective, the family 
Chilinidae is among the most primitive pulmo- 


*Corresponding author: dieguty@fcnym.unlp.edu.ar 


321 


nate gasteropods, and one of the first that 
conquered the aquatic environment. This is 
reflected in such primitive characters as a 
streptoneurous nervous system, horizontal 
lamellar tentacles, non-contractile pneumo- 
stome, and incomplete division of male and 
female ducts (Hubendick, 1947, 1978; 
Duncan, 1960a; Harry, 1964; Brace, 1983). 
More recently, based on analyses of 28S rRNA 
sequences, Dayrat et al. (2001) confirmed the 
monophyly of Hygrophyla, including the Chilini- 
dae at the base of this clade. 

The genus Chilina has the following distinc- 
tive characteristics: Shell: oval, oblong to ven- 
tricose, with last whorl expanded, spire erect 
or immersed, whorls generally convex (some- 
times carinate or angulose), aperture oval with 
columellar and parietal margins callous, with 
up to two oblique, tooth-like folds. Perio- 
stracum always present, olive-yellow, with dark 
brown longitudinal zigzag bands. Digestive 
system: Salivary glands slightly lobed; esopha- 
gus rather narrow; muscular stomach forming 
a caecum; anus opening to large mantle lobe; 
radula with numerous V-shaped tooth rows, 
with one central tooth and numerous lateral 
and marginal teeth laterally and posteriorly to 
central tooth. Reproductive system: genital 
system that is primitive among the Basomma- 
tophora because of the incomplete separation 


322 GUTIERREZ & RUMI 


TABLE 1. Number of specimens collected during different months at each rapid where Chilina 
iguazuensis was recorded, at Iguazú National Park, Argentina. 


Nandü Tacuara Mbigüa Leön Apepü Irene 
25°42'S, 29 36S: Zo S;, 257869, 29539 9, 2519601, 
54°25’W 54°21'W 54°26'W 54°14’°W 54°17°W 5423 W 
MLP 12526-28 12529-30 12532 12534 12533 12531 
Feb. 2004 70 
June 2004 20 
Sept. 2004 29 
Feb. 2005 7 10 19 16 
June 2005 20 
Dec. 2005 65 


of male and female reproductive ducts, ab- 
sence of fertilization chamber, and presence 
of very numerous penial spines; accessory 
seminal receptacle and calcareous granules 
in the vaginal lumen (Haeckel, 1911; Duncan 
1960a, b; Harry 1964; Castellanos & Gaillard, 
1981; Miquel, 1984). 

Here we describe a new species of this fam- 
ily, Chilina iguazuensis, from Iguazú National 
Park, Argentina; we provide descriptions of the 
shell and soft body parts, in particular the 
radula and internal organs. 


MATERIALS AND METHODS 


Specimens were collected during 2004 and 
2005 at Iguazu National Park, Misiones Prov- 


ince, Argentina (Fig. 1, Table 1). Soft parts 
were separated from the shell for subsequent 
processing, after relaxation in 10% Nembutal 
solution for 12 h, and fixed in modified Raillet- 
Henry solution for freshwater animals — 93% 
distilled water, 2% glacial acetic acid, 5% form- 
aldehyde, and 6 g sodium chloride per liter. 
Radulae were separated from the buccal mass 
and cleaned with sodium hypochlorite (Clorox). 

Six shell measurements were taken (Fig. 2): 
total length (TL), last whorl length (LWL), ap- 
erture length (AL), total width (TW), aperture 
width (AW), aperture projection (AP) and num- 
ber of teeth, following Gutiérrez et al. (1994) 
and Martin (2003). Radulae (n = 6) were ob- 
served under a scanning electron microscope 
at Museo de La Plata, Facultad de Ciencias 
Naturales y Museo, Universidad Nacional de 


FIG. 1. Rapids where Chilina iguazuensis was recorded in Iguazu River, within Iguazu National 


~ 


r 


Park, Argentina. A: Apepu; |: Irene; L: León; М: Mbigüa; N: Nandu; T: Tacuara. 


CHILINA IGUAZUENSIS N. SP. 323 


SNS Lits 
= 


FIG. 2. Shell measurements used for Chilina 
iguazuensis: TL: total length; LWL: last whorl 
length; AL: aperture length; TW: total width; AW: 
aperture width; AP: aperture projection. 


La Plata. Internal anatomy (n = 11) was ana- 
lyzed using a stereoscopic binocular micro- 
scope Leica MZ6 with camera lucida. 

Shell characters were compared to the origi- 
nal description of Chilina megastoma Hylton 
Scott, 1958, a species that also occurs in 
Iguazu National Park, but inhabits waterfalls 
of the Iguazu River basin. Anatomical traits 
were compared with the work of Ituarte (1997) 
about the latter species, as well as with our 
own measurements from 11 C. megastoma 
specimens collected at Arrechea Fall, Iguazu 
National Park, prepared following the above 
methodology. 

Average measurements of soft parts were 
divided by the length of the last whorl, in order 
to obtain size-free variables that facilitate com- 
parisons among different individuals and spe- 
cies. Length of last whorl was preferred over 
dimensions of soft parts because it was the 
most consistent measurement among individu- 
als, and it was not affected by relaxation tech- 
niques. 


FIG. 3. Shell of Chilina iguazuensis, n. sp. A, B: Adult specimen (Holotype) 
(18.08 mm); С, D: Juvenile specimen (6.27 mm). A, С = apertural view; В, D = 
dorsal view. 


324 GUTIERREZ & RUMI 


RESULTS 


Chilina iguazuensis, sp. nov. 
(Figs. 1-7) 


Type Locality 


Upper Iguazu River, Iguazu National Park, 
Misiones Province, Argentina. 


Type Material 


Holotype (MLP: 12526) and paratypes (MLP: 
12527, 12528) at Museo de La Plata; 
paratypes (MACN-In 37175) at Museo 
Argentino de Ciencias Naturales “Bernardino 
Rivadavia’. 


Etymology 


The specific name refers to the type locality, 
Iguazú River in Iguazu National Park. 


Diagnosis 


Shell globular, spire immersed in adult speci- 
mens (visible only in specimens less than 7 
mm long), length of aperture equal to length 
of last whorl; interior of shell markedly irides- 
cent. Radula: 57-65 teeth rows; 51-63 teeth 
per half-row; central tooth asymmetrical, bi- 
cuspid, both cusps serrated and second lat- 
eral tooth tetracuspid. Reproductive system: 
prepuce length 60% of penis sheath length; 
secondary bursa copulatrix tubular, short. 
Nervous system: mean distance between left 
pleural and parietal ganglia 1.39 mm (SD: 
0.97), approximately 7% of last whorl length. 


Description 


Shell (Fig. 3): Shell strong, oval, globular. Color 
purplish. Spire immersed, except in very 
small specimens (< 7mm). Last whorl very 
large, dilated, length equal to aperture 


FIG. 4. Radula of Chilina iguazuensis. A: General view; B: Lateral view; C: Central tooth and first 
lateral teeth; D: marginal teeth. Scale bars = 10 um. 


CHILINA IGUAZUENSIS N. SP. 325 


TABLE 2. Shell measurements of Chilina iguazuensis. All measurements in mm. Sp: specimen; TL: 
total length; LWL: last whorl length; AL: aperture length; TW: total width; AW: aperture width; AP: 


aperture projection. 


Date Sp. Ты LWL AL TW AW AP 

Nandü 

Holotype MLP12526 Feb. 2004 3.18.08 18.09 710.09 1487" 1245 ¥.14 
Paratype MLP12527 Feb. 2004 т AeA Ze!) 24 2-17-90 "1546" 620 
Paratype MLP12527 Feb. 2004 2 1602 76:62 76629 18.60 1.00 742 
Paratype MLP12527 Feb. 2004 A 18.397 10.39 “16.39 №22 "1176 - 6.96 
Paratype MLP12527 Feb. 2004 5.150 15.05" 45.05 1246 10.25 . 6:06 
Paratype MLP12527 Feb. 2004 O "Boo" Mos “Teo 183.37 "10.10 16:91 
Paratype MLP12528 Sept. 2004 1* .20,24- 202272024 16.70 1165: 648 
Paratype MLP 12528 Sept. 2004 2 120650" 20:66 ‘20667 18.82% 1345 8:99 
Paratype MLP 12528 Sept. 2004 3. 12240) 22.10. 2216 18.37 "18,064 “908 
Paratype MLP12528 Sept. 2004 4:19.70 + 78.76 2 19.76 1784 1236 - "8:07 
Paratype MLP12528 Sept. 2004 De РР 192837 199 727 
Paratype MLP12528 Sept. 2004 O Wo Tos “Wes “4146 1115 “694 
Paratype MLP 12528 Sept. 2004 FARO RON AO Zr 1.90 
Tacuara 

MLP 12529 June 2004 a GeO soe о 19.60 “17.15 1255 - 588 
MLP 12529 June 2004 2, AMAS 1228 ~ O96 8.09 Ot 
MLP 12529 June 2004 Sean 1222 1222 |. 9:91 1.37 “4.00 
MLP 12529 June 2004 a7 ZST 238901. 2910 27,22 "1410 OD 
MLP 12529 June 2004 Bee oe нае 
MLP 12529 June 2004 O. uaa. ta 2412 2210 1149 12.50. 003 


length. Well-defined growth lines crossed by 
spiral grooves, forming reticulate shell orna- 
mentation. A zigzag colored band on last 
whorl of juveniles, absent in adults. Aperture 
large, peristome outline oval. Labrum simple. 
Columellar callosity white, straight. One col- 
umellar tooth. Interior markedly iridescent. 
Parietal callosity weak, with soft white col- 
oration. Two whorls. Maximum size recorded 
24.12 mm total length. Table 2 lists shell 


measurements for 19 specimens. 


row based on counts of first rows (4-9). 
Mean number of teeth per half-row (not in- 
cluding central tooth) 51, ranging from 43 
(12.13 mm specimen) to 63 (22.95 mm 
specimen). Central tooth (Fig. 4C) asym- 
metrical, bicuspid, with one cusp slightly dis- 
placed to right; this latter cusp bifurcated in 
some specimens. External margins of both 
cusps serrate. Free part of central tooth not 
flat but with anteroposterior groove originat- 
ing from inward folding of basal plate, be- 


Digestive System: General morphology follows 
the basic plan described above for the ge- 
nus. In this section we provide a description 
of radular morphology. 

Radula with numerous tooth rows arranged 
in V-shape, as in all species of the family 
(Fig. 4A). All teeth except central united to 
membrane by base and an anterior projec- 
tion, base directed forward of radula and 
projection backward, and both situated at 
base of cusps (Fig. 4B). Six individuals rang- 
ing between 12.13 and 22.95 mm total length 
examined. Mean estimated number of rows 
58; number of rows 57 (12.13 mm specimen) 
to 65 (19.2 mm). Number of teeth per half- 


coming narrower and deeper posteriorly (Fig. 
4C). Aprotuberance at each margin of ante- 
rior area of tooth base. 

First lateral tooth tricuspid, with larger dag- 
ger-shaped mesocone higher and broader 
than other cusps (Fig. 4C). Ectocone of left 
tooth bifurcated in one individual. First lat- 
eral tooth slightly curved toward central tooth. 
Second lateral tooth tetracuspid, with smaller, 
divided ectocone (fourth cusp) and undivided 
endocone and mesocone. Contiguous teeth 
with more rounded appearance due to 
smaller cusps. Last marginal teeth simple, 
with poorly developed cusps, and ectocone 
divided into three minor cusps (Fig. 4D). 


326 GUTIERREZ & RUMI 


+ u ` 
Y en A } > 
/ + $e y. y 
I б 7 
| dg / 
| > ( — ES cou 
a 
\ 7 ` 
ls e ) 
© 
f 
/ | 
/ 
| а 
N o 
N 


9 
pra 


FIG. 5. Diagram of reproductive system of Chilina iguazuensis: ag: albumen gland; vd: 
vas deferens; dg: digestive gland; bc: bursa copulatrix; bcd: bursa copulatrix duct; fp: 
female pore; hd: hermaphrodite duct; mp: male pore; o: ovotestis; pp: prepuce; pr: pros- 
tate; ps: penis sheath; u: uterus; v: vagina; 2bc: secondary bursa copulatrix. Dotted line: 


body wall. Scale bar = 2 cm. 


Radular Formula: [51/(3-5) + 1/2] 57-65 [= 
number of right and left teeth/(number of 
cusps) + number of central teeth/number of 
cusps] extreme numbers for transversal rows. 

Reproductive System (Fig. 5): Ovotestis (her- 
maphroditic gonad) and a common duct an- 
terior to the separation of both systems. 
Female and male reproductive elements are 
described separately. Distinctive character- 
istics of C. iguazuensis: 


Female Genital System: Bursa copulatrix duct 


emerging from anterodorsal vagina and ex- 
tending alongside intestine, then passing 
under vagina and over uterus-vagina com- 
plex toward visceral mass; finally opening 
into flat oval bursa copulatrix next to ven- 
tricle. Secondary bursa copulatrix arising at 
base of uterus (oviduct); short (21% of length 
of the bursa copulatrix duct) and cylindrical, 
not expanded distally. 


CHILINA IGUAZUENSIS М. SP. ей 


FIG. 6. Diagram of nervous system of Chilina 
iguazuensis. Abbreviations for ganglia: Ic: left 
cerebral; Ipe: left pedal; Ip: left parietal; Ipl: left 
pleural; rc: right cerebral; rpe: right pedal; rp: right 
parietal; rpl: right pleural; si: subintestinal; v: vis- 
ceral. Scale bar = 1 mm. 


Male Genital System: Vas deferens emerg- 
ing from prostate; prostrate surrounding al- 
bumen gland (obvious in recently fixed 
specimens due to yellowish coloration). Vas 
passing under uterus and vagina, twisting 
upon itself, and finally located over vagina, 
once past final bend of the latter (Fig. 5). 
Vas deferens running laterally to buccal 
mass and not above vagina, coiling upon 
itself at this sector and making four loops; 
then entering body wall on right side with 


several minor turns and twists; this section 
twice as long as distance between vaginal 
and male pores. Vas deferens emerging 
from body wall at penial complex level and 
extending toward latter, crossing above pre- 
puce. At this section vas extending along- 
side penis sheath in straight trajectory, and 
entering the latter near buccal mass. Pre- 
puce length 60% of penis sheath length. 
Structure of prepuce, penis and penis sheath 
similar to those described by ltuarte (1997) 
for Chilina megastoma. 


Nervous System (Figs. 6, 7; Table 3): Pedal 


and cerebral ganglia connected by commis- 
sures and connectives, forming anterior 
nerve ring located at anterior half of buccal 
mass, only slightly posterior to beginning of 
esophagus. Long connective (ratio: 13.53 
of length of the last whorl) linking right pleu- 
ral ganglion to right parietal ganglion. Right 
parietal ganglion giving rise to two nerves, 
one to osphradium and one long, very thin 
connective to visceral ganglion behind pos- 
terior nerve ring. Two connectives on left 
side of anterior nerve ring, longer than those 
on right side, connecting left cerebral and 
pedal ganglia to pleural ganglion. Small con- 
nective (ratio: 4.79 of last whorl length) from 
pleural ganglion to left of parietal ganglion; 
this connective 65% shorter than right-side 
counterpart. Long connective (ratio: 18.43 
of last whorl length) linking left parietal gan- 
glion to subintestinal ganglion, located 
above posterior half of columellar muscle. 
One very short connective (ratio: 6.28 of last 
whorl length) linking subintestinal ganglion 
to visceral ganglion and closing posterior 
nerve ring. Two nerves arising from visceral 
ganglion toward visceral mass. One large 
nerve extending to right from subintestinal 
ganglion through columellar muscle to in- 
nervate distal-most vagina and accesory 
pneumostome. Pleurovisceral connectives 
with incomplete torsion characteristic of the 
genus. Table 3 shows proportions between 
length of connectives and length of last 
whorl. 


Distribution 


The new species was detected only at the 


type locality, Iguazú National Park, in the rap- 
ids of Upper Iguazu River listed in Table 1. 


328 GUTIERREZ & RUMI 


FIG. 7. Nervous system of Chilina iguazuensis in situ. 


DISCUSSION 


The species closest to C. iguazuensis 
based on shell, anatomical and distributional 
characteristics is Chilina megastoma (Fig. 8). 
The populations of C. iguazuensis were re- 
corded along the course of the upper Iguazu 
River, mainly in rapids. In contrast, C. mega- 
stoma only occurs in wet zones formed by the 
spray of the Iguazu River in the vicinity of 
Iguazú National Park, Argentina-Brazil. 

So far, two other Chilina species are known 
for areas geographically close to the range of 
the new species: Chilina gallardoi Castellanos 
& Gaillard, 1981, from Uruguay River, which 
is not connected to the Iguazu River; and 


Chilina guaraniana Castellanos & Miquel, 
1980, from the Parana River. The latter spe- 
cies was described based on materials col- 
lected in 1935 and not recorded since. The 
populations of both these species are located 
about 300 km south of Iguazu National Park. 
The internal anatomy of these two species is 
as yet unknown. Both are markedly different 
from C. iguazuensis and C. megastoma in 
shell morphology, with the aperture not greatly 
developed in either C. gallardoi or C. 
guaraniana. In addition, the aperture of C. 
gallardoi has two well-developed (columellar 
and parietal) teeth, and C. guaraniana has a 
single columellar tooth in a markedly angular 
aperture. 


CHILINA IGUAZUENSIS N. SP. 329 


FIG. 8. Shell of Chilina megastoma from Arrechea 
Falls, Iguazú National Park (TL = 13.3 mm). 


Another species with close geographic dis- 
tribution, Chilina parva Martens, 1868, inhab- 
its the states of Santa Catarina and Rio Grande 
do Sul, Brazil (Simone, 2006). This species is 
much smaller (5-8mm), with slight spire de- 
velopment and only one columellar tooth 
(Simone, 2006). 

Chilina megastoma was compared with the 
new species because its internal anatomy is 
known, and its geographical distribution is clos- 
est to that of C. iguazuensis. With respect to 
shell characters, aperture length is not equal 
to length of last whorl in C. megastoma, 
whereas C. iguazuensis has the same length; 
C. megastoma has two teeth on the labium and 
greater spire development, and it lacks inte- 
rior iridescence, whereas C. iguazuensis has 
just one teeth in its labium, the spire is im- 
mersed, and it is iridescent within. It is similar 
to the new species in the possession of greatly 
developed aperture and reticulate shell. Maxi- 
mum recorded size is 24.12 mm for C. 
iguazuensis, 17.7 mm for C. megastoma. 

Regarding anatomical traits, the two species 
differ in radular characteristics. Chilina 
iguazuensis has 58 tooth rows, with 51 teeth 
per half-row; the central tooth has serrated 
cusps, and the second lateral tooth is 
tetracuspid; C. megastoma has approximately 


40 tooth rows with 42 teeth per half-row, the 
central tooth lacks serrated cusps, and the sec- 
ond lateral tooth is tricuspid. 

Concerning the female reproductive system, 
the secondary bursa copulatrix of C. 
iguazuensis is uniformly wide along its entire 
length, whereas it expands terminally in C. 
megastoma. The course of the bursa copulatrix 
duct is similar in both species. Concerning the 
male reproductive system, prepuce length rep- 
resents 60% of the length of the penis sheath 
in C. iguazuensis, whereas it is 30 to 50% of 
penis sheath length in C. megastoma. 

The nervous system shows the pattern de- 
scribed by Ituarte (1997) for С. megastoma 
regarding the number of connectives, but with 
differences in length. The connectives of C. 
megastoma are generally longer, relative to 
length of last whorl, than in C. iguazuensis, 
except for the left pleuroparietal connective 
(Table 3). According to ltuarte (1997), in C. 
megastoma the length of the left cerebro- 
pleural connective is similar to that of the 
pleuroparietal connectives. However, our own 
dissections of relaxed specimens of this spe- 
cies indicate that the length of the left 
cerebropleural connective (11.29 ratio) is 
slightly more than twice that of the left 
pleuroparietal connectives (5.27 ratio). The left 
pleuroparietal and cerebropleural connectives 
are similar in C. iguazuensis. According to 
Ituarte (1997), the left parietal-subintestinal 
connective of C. megastoma is approximately 
three times as long as the pleuroparietal con- 
nective, but our own dissections indicate a 4.4 
ratio for these structures. This proportion is 
lower in C. iguazuensis (2.8 ratio). In addi- 
tion, a nerve arising at approximately two- 
thirds of the length of the left parietal- 
subintestinal connective, which was mentioned 
by Ituarte (1997) and observed in our dissec- 
tions of C. megastoma, was not detected in 
C. iguazuensis. 


ACKNOWLEDGEMENTS 


This study was financially supported by 
Consejo Nacional de Investigaciones 
Científicas y Técnicas (CONICET) (PIP 2711) 
and the Facultad de Ciencias Naturales y 
Museo, Universidad Nacional de La Plata 
(PN470). The authors wish to thank the staff 
at Centro de Investigaciones Ecológicas 
Subtropicales (CIES) in Iguazú National Park, 
and V. Núñez and N. Ferrando for their sup- 
port during field work. 


330 GUTIERREZ & RUMI 


LITERATURE CITED 


BRACE, R. C., 1983, Observations on the mor- 
phology and behaviour of Chilina fluctuosa 
Gray (Chilinidae), with a discussion on the early 
evolution of pulmonate gastropods. Philosophi- 
cal Transaction of the Royal Society of Lon- 
don, (ser. B), 300: 463-491 

CASTELLANOS, Z. A. de & M. C. GAILLARD, 
1981, Mollusca Gasteröpoda: Chilinidae. 
Fauna de Agua Dulce de la Republica Argen- 
tina. PROFADU (CONICET), Buenos Aires, 
15(4): 23-51. 

CASTELLANOS, Z. A. de & N. A. LANDONI, 
1995, Mollusca Pelecypoda y Gastropoda. Pp. 
759-801, in: E. C. LOPRETTO & G. TELL, eds., 
Ecosistemas de aguas continentales, meto- 
dologias para su Estudio, Vol. 2. Ediciones Sur, 
La Plata, Argentina. 3 vols., xxx + 1,401 pp. 

CASTELLANOS, Z. A. de & S. E. MIQUEL, 1980, 
Notas complementarias al género Chilina Gray 
(Mollusca Pulmonata). Neotröpica, 26: 171- 
178. 

CASTELLANOS, Z. A. de &S. E. MIQUEL, 1991, 
Distribución de los Pulmonata Basommato- 
phora. Fauna de Agua Dulce de la República 
Argentina. PROFADU (CONICET), Buenos 
Aires, 15(9): 1-9. 

DAYRAT, B., A. TILLIER, С. LECOINTRE & 5. 
TILLIER, 2001, New clades of euthyneuran 
gastropods (Mollusca) from 28S rRna Se- 
quences. Molecular Phylogenetics and Evolu- 
tion, 19: 225-235. 

DUNCAN, C. J., 1960a, The evolution of the 
pulmonate genital systems. Proceedings of the 
Zoological Society of London, 134: 601-609. 

DUNCAN, C. J., 1960b, The genital systems of 
the freshwater Basommatophora. Proceedings 
of the Zoological Society of London, 135: 339- 
ЗУБ: 

DUNCAN, С. J., 1975, Reproduction. Pp. 309— 
365, in: V. FRETTER & J. PEAKE, eds., Pulmo- 
nates, Vol. 1. Functional anatomy and 
physiology. London, Academic Press. xxix + 


Pp. 

GUTIERREZ, A., G PERERA, M. YONG & J. 
SANCHEZ, 1994, Estudio morfométrico en dos 
poblaciones del género Pomacea (Proso- 
branchia: Ampullariidae) de Cuba. Walkerana, 
7: 15-22. 

HAECKEL, W., 1911, Beiträge zur Anatomie der 
Gattung Chilina. Zoologische Jahrbücher, 
Supplement, 13: 89-136. 

HARRY, W. H., 1964, The anatomy of Chilina 
fluctuosa Gray reexamined, with prolegomena 


on the phylogeny of the higher limnic 
Basommatophora (Gastropoda: Pulmonata). 
Malacologia, 1: 355-385. 

HIDALGO, J. G., 1880, Description d’une 
nouvelle espece de Chilina. Journal de 
Conchyliologie, 20: 322-323. 

HUBENDICK, B., 1947, Phylogenetic relations 
between the higher limnic Basommatophora. 
Zoologiska Bidrag Fran Uppsala, 25: 141-162. 

HUBENDICK, B., 1978, Systematic and com- 
parative morphology of the Basommatophora. 
Pp. 1-47, in: V. FRETTER, & J. PEAKE, eds. Pul- 
monates, vol. 2A, Systematics, evolution, and 
ecology. London, Academic Press. xi + 540 pp. 

HYLTON SCOTT, M.I., 1958, Nueva especie de 
Chilina del norte Argentino. Neotropica, 4: 26- 
27. 

ITUARTE, С. F., 1997, Chilina megastoma Hylton 
Scott, 1958 (Pulmonata: Basommatophora): a 
study on topotypic specimens. American Ma- 
lacological Bulletin, 14: 9-15. 

MARTIN, P. R., 2003, Allometric growth an inter- 
population morphological variation of the fresh- 
water snail Chilina parchappii (Gastropoda: 
Chilinidae) in the Naposta Grande stream, 
southern Pampas, Argentina. Studies in Neo- 
tropical Fauna and Environment, 38: 71-78. 

MARSHALL, W. B., 1924, New species of mol- 
lusks of the genus Chilina. Proceedings of the 
United States National Museum, 66: 1-5. 

MARSHALL, W. B., 1933, New fresh-water gas- 
tropod mollusks of the genus Chilina of South 
America. Proceedings of the United States 
National Museum, 82: 1-6. 

MIQUEL, S. E., 1984, Contribuciôn al cono- 
cimiento biolögico de gasteröpodos pulmonados 
del area rioplatense, con especial referencia a 
Chilina fluminea (Maton). Tesis doctoral, 
Facultad de Ciencias Naturales y Museo, 
Universidad Nacional de la Plata. 133 pp. 

MIQUEL, $. E., 1987, Estudio micro-anatómico 
del complejo peniano en especies del genero 
Chilina Gray, 1828 (Gastropoda Basommato- 
phora). Notas del Museo de La Plata, 21 
(Zoologia 209): 131-142. 

SIMONE, L. R. L., 2006, Land and freshwater 
mollusc of Brazil. EGB, Fapesp., Sao Paulo. 
390 pp. 

VALDOVINOS, C. & J. STUARDO, 1995, Morfo- 
logia funcional de Chilina angusta (Philippi, 
1860), y evolución de Chilinidae. Resumos, II 
Congreso Latino-Americano de Malacología, 
Porto Alegre, Brasil: 43 


Revised ms. accepted 17 December 2007 


MALACOLOGIA, 2008, 50(1-2): 331-339 


CONVERGENCE CAUSED CONFUSION: ON THE SYSTEMATICS OF THE 
FRESHWATER GASTROPOD SULCOSPIRA PISUM (BROT, 1868) 
(CERITHIOIDEA, PACHYCHILIDAE) 


Frank Kóhler?**, Nora Brinkmann? € Matthias Glaubrecht' 


ABSTRACT 


We document a case of shell convergence in sympatric freshwater cerithioidean gastro- 
pods that has caused confusion in traditional classifications emphasizing shell features. 
Based on the comparative study of the operculum, radula, and embryonic shell obtained 
from dry remains of soft bodies, we present evidence that “Melania” pisum is not а thiarid 
species closely related to Balanocochlis glans (Busch, 1842), as has been supposed with 
respect to the very similar shells of both species. The species is transferred to the family 
Pachychilidae, because it shows various typical character states for the family. It is tenta- 
tively placed within the genus Sulcospira, which is endemic to Java. We assume that a 
similar shell shape has evolved in both species of not closely related gastropods through 
convergence, which once more reveals that purely shell-based classifications are particu- 


larly problematic. 


Key words: Cerithioidea, Thiaridae, Balanocochlis, Java, molluscan shell, convergence. 


INTRODUCTION 


Molluscan shells have been particularly em- 
phasized in classical systematic treatments. 
Indeed, shells bear many features that are 
convenient for taxonomic purposes, especially 
at the species level. An important aspect is 
that shell features are accessible also from dry 
material, which still represents the majority of 
museum holdings. In addition, dry shells of- 
ten constitute taxonomically or historically im- 
portant lots, such as types. Consequently, the 
importance of shell features for the recogni- 
tion of taxa is also acknowledged in modern 
systematic works (e.g., Smith, 1981; Ridgway 
et al., 1998; Vermeij & Snyder, 2006), as well 
as in field guides. On the other hand, the shell 
is particularly prone to different kinds of envi- 
ronmental pressures (Vermeij & Covich, 1978; 
Reid, 1986, 1992; Warner, 1996; West & 
Cohen, 1996; Reed & Janzen, 1999), which 
may lead to the evolution of convergent traits, 
as shown even for relatively complex shell 
structures, such as the clausilial apparatus 
(Moorsel et al., 2000). 

Since the late 20" century, shell shape con- 
vergence in gastropods has increasingly been 


highlighted as causing considerable confusion 
in taxonomic and systematic studies as dem- 
onstrated by, for example, Davis (1979) show- 
ing convergences among pomatiopsid, 
hydrobiid, littorinid, and cerithioidean gastro- 
pods, West & Cohen (1996) and Strong & 
Glaubrecht (2003) for thalassoid paludomids 
in Lake Tanganyika, and by Albrecht et al. 
(2004) for patelliform basommatophorans. 
The problem inherent to the traditional ap- 
proach of studying exclusively dry shells is that 
no other sources of information are used to 
evaluate the diagnostic value of shell features 
regarded as being taxonomically informative. 
Surprisingly, not a professional scientist, but 
rather the poet Edgar Allen Poe (1809-1849) 
was among the first (if not the first) to recog- 
nize that a reliable determination and classifi- 
cation of molluscs requires a combined analysis 
of the shell and soft body anatomy (Poe, 1839). 
In his scholarly textbook, he also distinguished 
— one of the first to do so — between the terms 
conchology (the study of shells) and malacol- 
ogy (the study of molluscs). Unfortunately, Poe’s 
insights were ignored for more than a century 
before first Keen (1936) and later Gould (1995) 
took notice of it. The study of Davis (1979) may 


‘Museum für Naturkunde, Humboldt-Universitat, Invalidenstrasse 43, 10115 Berlin, Germany 
“Present address: Cell Biology and Comparative Zoology, Department of Biology, Kgbenhavns Universitet, 


Universitetsparken 15, 2100 Copenhagen, Denmark 
“Corresponding author: frank.koehler@rz.hu-berlin.de 


332 KOHLER ETAL. 


be taken as a prime example of modern mala- 
cology, which seeks to combine detailed quan- 
titative and qualitative anatomy, multivariate 
analyses and shell morphology to untangle 
problems confounded in earlier works by 
convergences in hydrobioid snails. 

Also, in the polymorphic freshwater cerithioi- 
dean gastropods formerly subsumed under the 
term “melanians”, for a long time shell features 
were almost exclusively used for the classifi- 
cation of supraspecific taxa. For example, Brot 
(1874) stated that a siphonate aperture was 
typical for the genus Acrostoma Brot, 1870 
(synonym of the pachychilid genus Paracrosto- 
ma Cossmann, 1900; see Kohler & Glau- 
brecht, 2002). Later, Solem (1966) and Brandt 
(1974) suggested that Paracrostoma can be 
distinguished from the closely related genus 
Brotia H. Adams, 1866, by its generally coni- 
cal shell. However, recent revisions demon- 
strated that in Pachychilidae there is no 
correlation between shell shape and other mor- 
phological characters (Köhler & Glaubrecht, 
2001, 2003, 2005, 2006, 2007). Molecular and 
morphological evidence showed that similar 
shell forms (i.e., conical ones) reflect similar 
ecological adaptations rather than close rela- 
tionship (Glaubrecht & Köhler, 2004). By con- 
trast, it was shown that pachychilid species 
formerly treated as conspecific based on the 
possession of a similar shell can be differenti- 
ated by means of other morphological char- 
acteristics (e.g., Köhler & Glaubrecht, 2006). 
For a striking example of the systematic confu- 
sion arising from a deviant appraisal of shell 
features compare the influential view of 
Rensch (1934), Benthem Jutting (1956), and 
Brandt (1974), who assumed that Brotia 
costula (Rafinesque, 1833) is a widely distrib- 
uted and morphologically plastic species, ver- 
sus the revised concept of Köhler & Glaubrecht 
(2006). The latter authors delineated B. cos- 
tula in a much more restricted way and re- 
moved numerous taxa with similar shells from 
its synonymy. With respect to both geographi- 
cal distribution and morphological plasticity, 
the revised taxonomy revealed that the ranges 
of Brotia species are generally much more re- 
stricted than formerly assumed. 

In the present paper, we address the system- 
atic affinities of an enigmatic species with a 
confusing taxonomic history. Described as 
Melania pisum Brot, 1868, from Java, it has 
attracted little attention mainly because of its 
sparse occurrence in museum collections 
worldwide. Melania pisum was originally con- 
sidered as a member of Acrostoma Brot, 1868 
(Brot, 1874) (= Paracrostoma), but ignored by 


most subsequent authors. In the few accounts 
availabe, M. pisum was generally treated as a 
member of the Thiaridae Troschel, 1857, and 
either retained as a valid species within the 
genus Balanocochlis Fischer, 1885 (Leschke, 
1914; Benthem Jutting, 1956), or considered 
as being conspecific with Balanocochlis glans 
(Busch, 1842) because of the similar shell 
(Kohler & Glaubrecht, 2002). All previous sys- 
tematic treatments were, however, based solely 
on shell features. Comparative data, including 
shell morphometry, radular and embryonic shell 
morphology, show that M. pisum is clearly dis- 
tinct form Balanocochlis glans. Furthermore, we 
present evidence that this species is not a 
thiarid, as formerly assumed, but member of 
the Pachychilidae, a family not recognized by 
most previous authors. Hence, although 
conchologically very similar to each other, the 
thiarid Balanocochlis glans and the pachychilid 
M. pisum — which is tentatively placed within 
the genus Sulcospira — are not even closely 
related. We assume that a similar shell shape 
has evolved in both species convergently, which 
once more reveals that purely shell-based clas- 
sifications are particularly problematic. 


MATERIALS AND METHODS 


The study is based on the examination of dry 
shell material, including type material, from the 
Museum of Natural History, London (ВММН), 
the Geowissenschaftliche Sammlung, Univer- 
sität Bremen (GSUG), the Muséum d'Histoire 
Naturelle, Genève (MHNG), the Natural His- 
tory Museum Naturalis, Leiden (RMNH), the 
Museum of Comparative Zoology, Cambridge, 
Mass. (MCZ), and the Museum für Naturkunde, 
Berlin (ZMB). 

Dimensions of adult shells were measured 
with a calliper precise to 0.1 mm using stan- 
dard parameters as detailed by, e.g., Köhler & 
Glaubrecht (2006). Shell dimensions (H — 
height, B — breadth, LA — length of aperture, 
WA — width of aperture, BW — height of body 
whorl, N — number of remaining whorls) were 
statistically analysed with the statistic software 
SPSS vs. 12.0. Remains of soft bodies found 
in dry shells were watered before opercula, 
radulae, and embryonic shells were removed 
and cleaned mechanically (opercula and em- 
bryonic shells) or by proteinase K digestion 
(radulae). Radulae and embryonic shells were 
mounted on specimens stubs using adhesive 
pads, coated with gold-palladium and examined 
using a Jeol FSM 6300 scanning electron mi- 
croscope. 


SULCOSPIRA PISUM 333 


SYSTEMATIC DESCRIPTIONS 
Pachychilidae Troschel, 1857 
Sulcospira Troschel, 1858 
Sulcospira pisum (Brot, 1868), new comb. 


Melania pisum Brot, 1868: 54—55, pl. 2, fig. 5; 
Brot, 1870: 323. 

Melania (Acrostoma) pisum — Brot, 1874: 18, 
pla figs: 

Melania (Balanocochlis) pisum — Leschke, 
1914: 251. 

Balanocochlis pisum — Benthem Jutting, 1956: 
384-385. 

Balanocochlis glans [partim] — Köhler & 
Glaubrecht, 2002: 124. 


Type Material: Lectotype, herein designated, 
and paralectotype MHNG (ex coll. Brot, “Java”). 
size of the lectotype [mm]: H = 14.2, B = 10.0, 
LA= 5.1, WA= 10.1, BW = 12.9, N = 2.5. 


Other Material: Four dried shells MNHG (ex 
coll. Brot, “Java, leg. Schepman”); 40 dried 
shells RMNH 106883-4 (“Java, ex coll. 
Junghuhn, Reg.Nr. 2204, det. Van Benthem 
Jutting”); nine dried shells MCZ 346541 (“Java”, 
leg. Schepman, ex coll. Junghuhn, ex RMNH). 


Taxonomic Remarks: The species was first 
described by Brot (1868) from material collected 
by Petit (Brot, 1874: 18). The shell depicted here 
in Figure 1A is designated as the lectotype of 
Melania pisum Brot, 1868, in order to preserve 
stability of nomenclature according to the stipu- 


FIG. 1. Shells (2x natural size). A: Lectotype of Melania pisum MNHG, Brot collection (Java); B: 
Paralectotype MNHG, Brot collection (Java); C: Three shells MNHG, Brot collection (Java); D: Three 
shells RMNH 106884 (Java); E: Syntype of Melania glans Busch, 1842 GSUG 14583; F: Balanocochlis 
glans ZMB 46.049 (SW Java, Palabuan); G: Balanocochlis glans ZMB 102.695 (New Ireland); H: 
Operculum of S. pisum RMNH 106884; |: Operculum of B. glans ZMB 46.049 (Java). Scale bar for 
shells = 10 mm. 


334 KOHLER ETAL. 


lations of § 74.7 of the Code of Zoological No- 
menclature (ICZN 1999). The paralectotype is 
depicted in Fig. 1B. There is a second lot in the 
Brot collection (MNHG) comprising four shells 
collected by Schepman as is evident from the 
label on the cardboard onto which the shells 
were mounted. Being of a deviant provenance, 
these shells are not considered as types. Later, 
Brot (1874) affiliated M. pisum with the genus 
Acrostoma Brot, 1870, and stated that a 
siphonate aperture is characteristic for this 
group. Because the name Acrostoma was pre- 
occupied, the valid name of the genus is Para- 
crostoma Cossmann, 1900 (Kôhler & 
Glaubrecht, 2002). Based on close concho- 
logical similarity with Balanocochlis glans, M. 
pisum was also considered a thiarid. Leschke 


(1914) and Benthem Jutting (1956) retained 
pisum as a valid species within Balanocochlis 
(to be distinguished from B. glans and B. glandi- 
formis by a more globular shape of the body 
whorl and generally smaller size), whereas 
Kohler & Glaubrecht (2002) suggested that it 
may be a junior synonym of B. glans. 


Shell Morphology (Fig. 1A—D): Comparatively 
small, with a globular shape. Spire always trun- 
cated with usually less than three remaining 
whorls. Body whorl large compared to rest of 
shell, accounting for around 90% of entire shell 
height, inflated, and convexly rounded in diam- 
eter. Scupture smooth except for growth lines. 
Colour chestnut brown, sometimes with darker 
vertical flames or lines. Aperture with thick col- 


FIG. 2. Radular and embryonic shell morphology. A-D: SEM images of a radular fragment of S. pisum 
extracted from dry shell material (RMNH 106884). A: View from above; B: Detail showing central and 
lateral teeth; C: Anterior viewed obliquely at 45° from above showing the shape of radular denticles; 
D: Detail showing marginal teeth; E-G: SEM images of a radular fragment of B. glans extracted from 
dry shell material (BMNH 2128; Solomon lls., Guadalcanal). E: View from above; Е: Detail showing 
central and lateral teeth; G: Detail showing lateral teeth; H: SEM images of embryonic shell of S. 
pisum (RMNH 106884), front view (above) and apical view (below). Scale bars = 100 um. 


SULCOSPIRA PISUM 335 


umellar margin, wide, laterally and terminally 
rounded, pointed above; lower margin being 
conspicuously protracted. Average size (n = 52, 
median and standard deviation in parentheses): 
H = 11.24 (1.57) mm, B = 8.32 (1.06) mm, LA= 
4.36 (0.66) mm, WA = 7.77 (0.96) mm, BW = 
10.46 (1.54) mm, N = 1.5 (0.61). 


Operculum: Ovate with almost a central 
nucleus, comprising four to five whorls (RMNH 
106883-4, various opercula of corresponding 
shape). 


Embryonic Shell Morphology (Fig. 2E): Brot 
(1874) found embryonic shells in one of the 
types comprising two whorls and exhibiting 
spiral colour bands. The whereabouts of these 
shells are unclear. We found three embryonic 
shells in the remains of one dried body (RMNH 
106884) with a height of about 1.2 mm com- 
prising 1.5 whorls. Sculpture is smooth, with 
faint growth lines. Apical whorl is inflated and 
dome-like. 


Radula (Fig. 2A-D): Two radula fragments 
isolated from dry remains of soft bodies 
(RMNH 106884) showed exactly the same 
morphology. The radula is taenioglossate, typi- 
cally pachychilid. Seen from above, central 
teeth are squarish; their cutting edge consists 
of a pronounced main cusp flanked by three 
accessory cusps that taper in size. Their up- 
per rim is convex, their lower rim is concave 
by the extending glabella. Marginal margins 
of the glabella are only slightly concave. Lat- 
eral teeth possess a triangular-shaped main 
cusp with two to three outer and one inner 
accessory cusp, their outer lateral margin sup- 
ports a lateral flange, ventral surface with con- 
spicuous glabella. Inner and outer marginals 
are hooked, each possesses a large, well 
rounded main denticle and a much smaller 
inner cusp. Outer marginals have a lateral 
flange at their exterior side. 


Soft Body Anatomy: Unknown for the lack of 
preserved material. The presence of embry- 
onic shells shows that S. pisum is a brooder. 


Distribution: All known material originates 
from Java; more precise locality information 
is not available from the historical museum 
material. Recent efforts by the authors and 
their collaborators to find the species in the 
field were not successful thus far. 


DISCUSSION 


Freshwater Cerithioidean Gastropods — Sketch- 
ing the Systematic Framework 


Tropical cerithioidean freshwater gastro- 
pods, currently known to represent several 
distinct freshwater radiations, were originally 
held to represent one large monophyletic 
group. This group was initially called the 
melanians (or Melaniidae), but later renamed 
Thiaridae because the generic name Thiara 
Rôding, 1798, has priority over Melania 
Lamarck, 1799 (reviewed in Glaubrecht, 1996, 
1999, 2006). Later recognized as polyphyletic, 
this group was progressively split into various 
distinct families, each of it characterized by 
typical radular, operculum, and soft body char- 
acteristics. These lineages are also recognized 
in molecular phylogenetic studies (Glaubrecht, 
1996, 1999, 2006; Lydeard et al., 2002; Köhler 
et al., 2004). This led to a more restricted defi- 
nition of the true Thiaridae and the delimita- 
tion of further freshwater lineages, such as the 
Pachychilidae, which were previously not 
widely accepted (reviewed in Köhler & 
Glaubrecht, 2002). These changing system- 
atic concepts and the inconsistent use of taxo- 
nomic names have caused considerable 
confusion. Today we know that the true 
Thiaridae and the Pachychilidae are only dis- 
tantly related to each other, representing lin- 
eages that colonized freshwater independently 
(Lydeard et al., 2002; Kohler et al., 2004). Their 
members are most easily recognized by 
means of their divergent — but within each fam- 
ily conserved — radula and operculum morphol- 
ogy (e.g., Köhler & Glaubrecht, 2001). 
Thiaridae generally possess a paucispiral 
operculum (Fig. 11), whereas Pachychilidae 
always have a multispiral operculum (Fig. 1H). 
In addition, the radula shows a variety of 
marked differences, such as varying shapes 
and relative sizes of teeth (compare e.g. Fig. 
2A-D with 2E, F). 


Diagnostic Value of Shell Features in Fresh- 
water Gastropods 


Contrary to Poe’s (1839) credo, in traditional 
classifications shell features were often exclu- 
sively used for the delineation of gastropod 
taxa. This holds true also for most freshwater 
cerithioideans, such as the species relevant 
in the context of the present study (B. glans, 


336 KOHLER ET AL. 


S. pisum). Their traditional placement within 
the Thiaridae merely reflects the unawareness 
of former workers that, in addition to the 
Thiaridae, there are further freshwater 
cerithioidean lineages while the affiliation of 
M. pisum with Balanocochlis by Leschke 
(1914) and Benthem Jutting (1956) solely rests 
on the postulate that a similar shell indicates 
close relationship (Fig. 1 compares these 
taxa). In fact, the shells of both species are 
similar in regard to their shape, colour, sculp- 
ture, and form of aperture. In illustrations of 
shells, Van Benthem Jutting (1956) showed 
the body whorl of pisum as being of a more 
globular shape compared to B. glans. How- 
ever, the examination of a large series of shells 
revealed that B. glans is quite variable and 
that globular shells also occur in this species 
(Brinkmann & Glaubrecht, unpubl. data). More 
reliable than with respect to overall shell shape, 
both taxa can be discriminated by means of 
shell size parameters. One-way ANOVA re- 
vealed that shells of S. pisum differ from those 
of B. glans in regard to various morphometric 
parameters (H, B, LA, WA, BW, N, H/B, H/LA, 
H/BW) with statistical significance at the 0.1% 
level (p < 0.000). Re-classification of shells by 
discriminant analysis (one discriminant func- 
tion, Wilk’s lamda = 0.213, p < 0.000) resulted 
in 100% correct reassignment of S. pisum 
shells (n = 52) and 99.4% correct reassign- 
ment of B. glans shells (n = 349), thus strongly 
corroborating the results of the ANOVA (dia- 
grammatically represented in Fig. 3). Thus, 
contrary to the assumption of Kohler & 
Glaubrecht (2002), our findings support the 
retention of S. pisum as a species distinct from 
B. glans, as suggested by Leschke (1914) and 
Benthem Jutting (1956). However, while the 
analyses of shell parameters demonstrate that 
close similarity does not imply identity in this 
case — the similar shells say nothing about the 
phylogenetic relationships of the species in- 
volved. Instead, we found that both species 
display marked morphological differences in 
the radula and operculum, which imply place- 
ment of each in distinct families (Thiaridae and 
Pachychilidae, respectively). Because the 
globular, almost limpet-like shape of shells is 
Unusual in both families of freshwater 
cerithioideans, we assume that it has evolved 
in parallel — possibly due to adaptation to simi- 
lar habitat conditions and lifestyles. Similar 
cases of homoplastic shell shapes have been 
reported from other Asian pachychilid species 
(Kohler & Glaubrecht, 2005, 2006), as well as 


further groups of freshwater gastropods (Davis, 
1979; West & Cohen, 1996; Strong & Glaub- 
recht, 2003; Albrecht et al., 2004). Hence, 
whether shell features can be con-sidered as 
being of diagnostic value with respect to the 
systematic treatment of a given taxon has to 
be tested in the context of more comprehen- 
sive analytical work. 


Systematic Implications 


As stated above, the former systematic 
placement of M. pisum in the Thiaridae merely 
reflects the traditional treatment of freshwater 
cerithioideans (e.g. Thiele, 1928, 1929-1935; 
Leschke, 1914; Benthem Jutting, 1956). As 
judged from the presence of typical charac- 
teristics of the operculum and radula, however, 
only B. glans is a genuine member of this fam- 
ily (paucispiral operculum with basal nucleus; 
radula with comparatively small central teeth, 
lateral teeth with a large lateral flange, and 
marginal teeth with numerous cusps; Figs. 1, 
2; see also Rensch, 1934; Benthem Jutting, 
1956; Starmühlner, 1976; Glaubrecht & 
Brinkmann, unpubl. data). By contrast, S. 
pisum exhibits characteristics typical only for 
Pachychilidae. These include possession of a 
multispiral operculum with a central nucleus 
and a radula with pronounced central teeth, 
lateral teeth with a short lateral flange, mar- 
ginal teeth with only two cusps. Also its em- 
bryonic shell closely resembles that of other 
Southeast Asian pachychilids (Köhler & 
Glaubrecht, 2001, 2005, 2007). Since pisum 
is evidently a pachychilid, it cannot be amem- 
ber of the thiarid genus Balanocochlis. To de- 
duce its correct generic placement within the 
Pachychilidae, however, is difficult. A molecu- 
lar phylogenetic analysis of the Asian 
Pachychilidae showed that the morphological 
characteristics found to be typical for the en- 
tire family (radula, operculum) are widely con- 
served among its representatives and 
therefore not suitable to infer the generic place- 
ment of the present species (Köhler & 
Glaubrecht, 2001, 2002; Kohler et al., 2004). 
Instead, it was demonstrated that the genera 
of SE Asian Pachychilidae (Adamietta, Brotia, 
Jagora, Paracrostoma, Sulcospira, Tylo- 
melania) essentially differ with respect to their 
reproductive anatomy (gonads, gonoducts, 
brooding structures) and, to a lesser degree 
in the embryonic shell morphology (Kôhler & 
Glaubrecht, 2001, 2003, 2005, 2006, 2007; 
Rintelen & Glaubrecht, 2005). 


SULCOSPIRA PISUM 


337 


Height of shell 


5 


10 


© Sulcospira pisum 


% Balanocochlis glans 


15 20 25 


Breadth of shell 


FIG. 3. Comparison of shells by means of the parameters 
height and breadth of the thiarid Balanocochlis glans (n = 154) 
and the pachychilid Sulcospira pisum (n = 52). 


Because the reproductive anatomy of pisum 
remains unknown due to the absence of pre- 
served material, it is not possible to reconstruct 
its generic affinities without ambiguity. A dome- 
like inflated embryonic shell with a smooth 
sculpture, as shown here for pisum, is known 
from a total of three pachychilid genera or 
groups: The so-called “Brotia testudinaria- 
group” (Köhler & Glaubrecht, 2001), the ge- 
nus Sulcospira endemic to Java (see Kohler 
& Glaubrecht, 2005), and Paracrostoma en- 
demic to India (Kohler & Glaubrecht, 2007). 
Because the “Brotia testudinaria-group” rep- 
resents an informal grouping only, which 
awaits critical revision, and Paracrostoma is 
restricted to southern India, Sulcospira is cur- 


rently rendered the only available generic 
name. Therefore, the pachychilid species 
pisum is here tentatively placed within this 
genus until a more detailed revision of all 
Javanan Pachychilidae becomes available. 


ACKNOWLEDGEMENTS 


We wish to thank two anonymous reviewers 
as well as George Davis for carefully reading 
the manuscript. Their suggestions and correc- 
tions help much to improve the quality of this 
paper. Jason Dunlop (Berlin) kindly helped to 
improve the style of the text, which is grate- 
fully acknowledged. 


338 KOHLER ET AL. 


LITERATURE CITED 


ALBRECHT, C., T. WILKE, K. KUHN & B. 
STREIT, 2004, Convergent evolution of shell 
shape in freshwater limpets: the African genus 
Burnupia. Zoological Journal of the Linnean 
Society, 140: 577-586. 

BENTHEM JUTTING, W. $. S. v., 1956, System- 
atic studies on the non-marine Mollusca of the 
Indo-Australian archipelago. V. Critical revision 
of the Javanese freshwater gastropods. 
Treubia, 23: 259-477. 

BRANDT, R.A. M., 1974, The non-marine aquatic 
Mollusca of Thailand. Archiv für Mollusken- 
kunde, 105: 1-423. 

BROT,A., 1868, Additions et corrections au cata- 
logue systématique des espéces qui 
composent la famille des mélaniens. Materiaux 
pour servir a Гешае de la famille des 
mélaniens, 2. Geneve, Chez George. ix + 64 
pp., 3 pls. 

BROT. A., 1870, Catalogue of the recent spe- 
cies of the family Melanidae. American Jour- 
nal of Conchyology, 6: 271-325. 

BROT, A., 1874-1879, Die Melaniaceen 
(Melanidae) in Abbildungen nach der Natur mit 
Beschreibungen. In: H. C. KUSTER, ed., Syste- 
matisches Conchylien-Cabinet von Martini und 
Chemnitz, 1(24): 488 pp., 49 pls. Nurnberg, 
Bauer & Raspe. 

DAVIS, G. M., 1979, The origin and evolution of 
the gastropod family Pomatiopsidae, with em- 
phasis on the Mekong River Triculinae. Mono- 
graphs of the Academy of Natural Sciences of 
Philadelphia, 20: 1-120. 

GLAUBRECHT, M., 1996, Evolutionsökologie 
und Systematik am Beispiel von Süß- und 
Brackwasserschnecken (Mollusca: Caeno- 
gastropoda: Cerithioidea): Ontogenese- 
Strategien, paläontologische Befunde und 
Historische Zoogeographie. Leiden, Backhuys 
Publishers. 544 pp. 

GLAUBRECHT, M., 1999, Systematics and the 
evolution of viviparity in tropical freshwater 
gastropods (Cerithioidea: Thiaridae sensu lato) 
— an overview. Courier Forschungs-Institut 
Senckenberg, 215: 91-96. 

GLAUBRECHT, M., 2006, Independent evolution 
of reproductive modes in viviparous freshwa- 
ter Cerithioidea (Gastropoda, Sorbeoconcha) 
— a brief review. Basteria, 69: Suppl. 3: 23-28. 

GLAUBRECHT, M. & F. KOHLER, 2004, Radiat- 
ing in a river: systematics, molecular genetics 
and morphological differentiation of viviparous 
freshwater gastropods endemic to the Kaek 
River, Central Thailand (Cerithioidea, Pachy- 
chilidae). Biological Journal of the Linnean 
Society of London, 82: 275-311. 

GOULD, S. J., 1995, Dinosaurs in a haystack. 
Reflections in natural history. New York, Har- 
mony Books/Crown Publishers. 496 pp. 

ICZN, 1999, International Code of Zoological 
Nomenclature, 4" edition. London, International 
Trust for Zoological Nomenclature. 306 pp. 

KEEN, A. M., 1939, Edgar Allan Poe‘s concho- 
logical text. The Nautilus, 50: 42-44. 


KOHLER, F. & M. GLAUBRECHT, 2001, Toward 
a systematic revision of the Southeast Asian 
freshwater gastropod Brotia H. Adams, 1866 
(Cerithioidea: Pachychilidae): an account of 
species from around the South China Sea. 
Journal of Molluscan Studies, 67: 281-318. 

KOHLER, F. & M. GLAUBRECHT, 2002, Anno- 
tated catalogue of the nominal taxa of South- 
east Asian freshwater gastropods, family 
Pachychilidae Troschel, 1857 (Mollusca: 
Caenogastropoda: Cerithioidea), with an evalu- 
ation of the types. Mitteilungen aus dem Mu- 
seum für Naturkunde Berlin, Zoologische 
Reihe, 78: 121-156. 

KOHLER, F. & M. GLAUBRECHT, 2003, Mor- 
phology, reproductive biology and molecular 
genetics of ovoviviparous freshwater gastro- 
pods (Cerithioidea: Pachychilidae) from the 
Philippines, with description of a new genus 
Jagora. Zoologica Scripta, 32: 35-59. 

KOHLER, Е. & M. GLAUBRECHT, 2005, Fallen 
into oblivion — the systematic affinities of the 
enigmatic Sulcospira Troschel, 1857 (Cerithio- 
idea: Pachychilidae), a genus of viviparous 
freshwater gastropods from Java. The Nauti- 
lus, 11: 15-27. 

KOHLER, F. & M. GLAUBRECHT, 2006, A sys- 
tematic revision of the Southeast Asian fresh- 
water gastropod Brotia (Cerithioidea: 
Pachychilidae). Malacologia, 48: 159-251. 

KOHLER, F. & M. GLAUBRECHT, 2007, Out of 
Asia and into India — On the molecular phylog- 
eny and biogeography of the endemic fresh- 
water gastropod Paracrostoma Cossmann, 
1900 (Caenogastropoda: Pachychilidae). Bio- 
logical Journal of the Linnean Society of Lon- 
don, 91: 627-651. 

KÔHLER, F., T. v. RINTELEN, A. MEYER & M. 
GLAUBRECHT, 2004, Multiple origin of vivi- 
parity in Southeast Asian gastropods (Cerithioi- 
dea: Pachychilidae) and its evolutionary 
implications. Evolution, 58: 2215-2226. 

LESCHKE, M., 1914, Zur Molluskenfauna von 
Java und Celebes. Mitteilungen aus dem 
Naturhistorischen Museum in Hamburg, 31: 
205-284. 

LY DEARD, С М E.YHOLZNAGEL ¿Mi 
GLAUBRECHT & W. F. PONDER, 2002, Mo- 
lecular phylogeny of a circum-global, diverse 
gastropod superfamily (Cerithioidea: Mollusca: 
Caenogastropoda): pushing the deepest phy- 
logenetic limits of mitochondrial LSU rDNA 
sequences. Molecular Phylogenetics and Evo- 
lution, 22: 399-406. 

MOORSEL, С. H. M. Е. С. M. DIJKSTRA & Е. 
GITTENBERGER, 2000, Molecular evidence 
for repetitive parallel evolution of shell struc- 
ture in Clausiliidae (Gastropoda, Pulmonata). 
Molecular Phylogenetics and Evolution, 17: 
200-208. 

POE, E.A., 1839, The conchologist’s first book: 
or, a system of testaceous malacology. Phila- 
delphia, Haswell, Barrington & Haswell. 156 
pps, 12215. 

REED, W. |. & Е. J. JANZEN, 1999, Natural se- 
lection by avian predators on size and colour 
of a freshwater snails (Pomacea flagellata). 


SULCOSPIRA PISUM 200 


Biological Journal of the Linnean Society, 67: 
331-342. 

REID, D. G., 1986, The littorinid molluscs of man- 
grove forests in the Indo-Pacific region. The 
genus Littoraria. London, British Museum 
(Natural History). 227 pp. 

REID, D. G., 1992, Predation by crabs on 
Littoraria species (Littorinidae) in a Queensland 
mangrove forest. Proceedings of the Third In- 
ternational Symposium on Littorinid Biology, 
141-151. 

RENSCH, B., 1934, SüRwassermollusken der 
deutschen limnologischen Sunda-Expedition. 
Archiv für Hydrobiologie, Suppl. 8: 203-254. 

RIDGWAY, T. M., B. A. STEWART, G. M. 
BRANCH &A. N. HODGSON, 1998, Morpho- 
logical and genetic differ-entiation of Patella 
granularis (Gastropoda: Patellidae): recogni- 
tion of two sibling species along the coast of 
southern Africa. Journal of Zoology, 245: 317- 
313. 

RINTELEN, T. v. & M. GLAUBRECHT, 2005, 
Anatomy of an adaptive radiation: a unique 
reproductive strategy in the endemic freshwa- 
ter gastropod Tylomelania (Cerithioidea: 
Pachychilidae) on Sulawesi, Indonesia, and its 
biogeographic implications. Biological Journal 
of the Linnean Society, 85: 513-542. 

SMITH, J. E., 1981, The natural history and tax- 
onomy of shell variation in the periwinkles 
Littorina saxatilis and Littorina rudis. Journal 
of the Marine Biological Association of the 
United Kingdom, 61: 215-241. 

SOLEM, A., 1966, Some non-marine Mollusks 
from Thailand, with notes on classification of 
the Helicarionidae. Spolia zoologica Musei 
Hauniensis, 24: 1-110. 


STARMUHLNER, F., 1976, Beiträge zur Kennt- 
nis der Süßwassergastropoden pazifischer In- 
seln. Ergebnisse der Österreichischen 
Indopazifik-Expedition des 1. Zoologischen In- 
stitutes der Universität Wien. Annalen des 
Naturhistorischen Museums in Wien, (B), 80: 
473-656. 

STRONG, Е. Е. & М. GLAUBRECHT, 2003, 
Anatomy and systematic affinity of Stanleya 
neritinoides (Smith, 1880), an enigmatic mem- 
ber of the thalassoid gastropod species flock 
from Lake Tanganyika, East Africa 
(Cerithioidea, Paludomidae). Acta Zoologica, 
84: 249-265. 

THIELE, J., 1928, Revision des Systems der 
Hydrobiiden und Melaniiden. Zoologische Jahr- 
bücher, Abteilung Systematik, 55: 351-402. 

THIELE, J., 1929-1935, Handbuch der Systema- 
tischen Weichtierkunde, 1-2. Jena, Gustav 
Fischer. 1,154 pp. 

VERMEIL, С. J. 8 A. P. COVICH, 1978, Coevo- 
lution of freshwater gastropods and their preda- 
tors. The American Naturalist, 112: 833-843. 

VERMEIL, С. J. & M. A. SNYDER, 2006, Shell 
characters and taxonomy of Latirus and related 
fasciolariid groups. Journal of Molluscan Stud- 
les, 72: 413-424. 

WARNER, G. F., 1996, Factors affecting the se- 
lection of pond snail prey by signal crayfish. 
Freshwater Crayfish, 11: 194—202. 

WEST, K. & A. COHEN, 1996, Shell microstruc- 
ture of gastropods from Lake Tanganyika, Af- 
rica: adaptation, convergent evolution, and 
escalation. Evolution, 50: 672-681. 


Revised ms. accepted 16 December 2007 


= E 


На рее ый 


[ALI mn MA o 
Bi 


7 ies 
ty i и 


1 
ais DE LT > 
Er 


Ad 


rere a 


A à a, 
| 


pos 


LE 


dE à 


er ie 
PE E mt 


И lent 
Bane 2008 we 


Ne {es 


ти a 


fost bande Res 


ee ii = AA 


+ Le E 
wi‘ 4 
Aiton “A 


Sg saint CE + Su er Bier 
Hg ite po A 


Frs ene 


hr 


ee de oe | bien 
Léon нар ABP ae rh aile 


e PAS 
oa ae ri Hs 
ARE 


4 e и ore! | 
er. u a Kane Era er 


a 5 


$ E CO ENGIN ut à 


pete a 
Bee tes 


Di Fe aie 
RR, 3 


7 ker + padre be 
Ah es aS: Lean de 


ER Be Meuse Аха & > a E he 


er as EUR cel MEN 


| nn Pag gis 
e de eee: 


i eons т. Е. RN 


$ ae 


EN 


PASEO: р. 
gt se №0 ah 


а ot phy 
¿FE 


MALACOLOGIA, 2008, 50(1-2): 341-345 


THE EASTERN MUDSNAIL, IEYANASSA OBSOLETA, ACTIVELY FORAGES FOR, 
CONSUMES, AND DIGESTS CYSTS OF THE DINOFLAGELLATE, 
SCRIPPSIELLA LACHRYMOSA 


Agneta Persson", Barry С. Smith", Mark S. Dixon! & Gary H. Wikfors' 


ABSTRACT 


The Eastern mudsnail, /lyanassa obsoleta, was attracted to, consumed, and digested 
resting cysts of the dinoflagellate Scrippsiella lachrymosa when cysts were presented in 
grazing experiments. Twenty snails were observed individually for one hour in petri dishes 
divided into four parts wherein cysts were present in one quadrant, sediment particles of 
the same size range were in another quadrant, and two quadrants were free of particles. 
Actively foraging snails were nearly twice as likely to be found in quadrants containing S. 
lachrymosa cysts as in the other quadrants until cysts were consumed. Microscope obser- 
vations of fecal pellets from snails feeding on cysts revealed digestive destruction of the 
cysts. These findings indicate that deposit-feeding grazers can actively seek dinoflagellate 
cysts as a food item, thereby influencing distribution of cysts and subsequent germination 


of dinoflagellate vegetative cells. 


Key words: deposit feeding, dinoflagellate resting stages, cyst ecology. 


INTRODUCTION 


Dinoflagellate cysts have, until recently, been 
largely regarded as inert particles, unaffected 
by processes other than sedimentation and 
accumulation. However, evidence is accumu- 
lating that many different animals ingest and 
digest, or at least destroy, dinoflagellate cysts 
when they are grazing (Persson & Rosenberg, 
2003). Cysts are frequently found in the fecal 
pellets of deposit feeders (Persson & 
Rosenberg, 2003; Kremp et al., 2003), cope- 
pods (Reid & Boalch, 1987), and filtering 
bivalves (Bravo et al., 1998, and references 
therein). Cysts are known to contain lipid and 
starch (Dale, 1983), and therefore could con- 
stitute a source of nutrition for grazers, but it 
is unclear if the cysts are eaten indiscriminately 
or if they are actively sought. In this experi- 
ment, we used the eastern mudsnail, //vanassa 
obsoleta, as a model deposit-feeding grazer 
to determine if cysts attract active grazing by 
a foraging, surface-deposit feeder. The cysts 
used were produced by pure cultures of 
Scrippsiella lachrymosa. 


MATERIALS AND METHODS 


The strain of Scrippsiella lachrymosa (strain 
B10) used was kindly provided by Anke Kremp, 
Woods Hole Oceanographic Institution. Cul- 
tures were grown in f/10-enriched Milford Har- 
bor seawater (a variation of f/2; Guillard & 
Ryther, 1962) with 5% sediment extract, and 
encystment was induced by transfer to f/2 with- 
out nitrogen enrichment (Smith & Persson, 
2004, provide details of encystment method). 
Cysts were sonicated 4 min (Sonicor SC-50) 
and rinsed thoroughly by spraying with filtered 
seawater on a 20 um sieve to remove any re- 
maining debris from the culturing medium. 

The /lyanassa obsoleta were collected at Fort 
Trumbull Beach, Milford, Connecticut, USA, 
from the intertidal zone on December 31, 2002, 
and kept in running seawater until the start of 
the experiment 17 days later. 

The experimental containers are shown in 
Figure 1. Twenty, 90 mm glass petri dishes, 
marked with a pencil on the outside of the 
bottom into four equal quadrants, were placed 
on a table, at room temperature, and filled with 


‘National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Northeast Fisheries Science Center, 


Milford Laboratory, Milford, Connecticut, 06460, U.S.A. 
“Smedjebacksvágen 13, SE-771 90 Ludvika, Sweden 
“Corresponding author: barry.smith@noaa.gov 


342 PERSSON ETAL. 


FIG. 1. Petri dish divided into quadrants with (1) cysts of the dinoflagel- 
late, Scrippsiella lachrymosa, (2) no particles, (3) sediment 20-100 um, 
and (4) no particles, used for selection experiment with Eastern mud 
snails, /lyanassa obsoleta. 


40 ml filtered seawater. In one quadrant of the 100 um) were placed. The other two quadrants 
dish, Scrippsiella lachrymosa cysts were placed of the dish were left empty of particles. Petri 
~800 cysts/dish), in the quadrant diagonal to dishes were oriented haphazardly so that pos- 
the cysts, sieved sediment particles from Fort sible, unknown factors of orientation in the room 
Trumbull Beach of the same size range (20- would not influence the results. 


FIG. 2. Photomicrograph of fecal pellet from an Eastern mud snail, 
llyanassa obsoleta, that had fed on cysts of the dinoflagellate, Scrippsiella 
lachrymosa. Partially and fully degraded material from cysts is present, 
indicating digestive degradation of the cysts. 


SNAILS EAT DINOFLAGELLATE CYSTS 


Cell means of transformed number of snails 


cysts empty 2 empty 4 sediment 
Treatment 


FIG. 3. Mean (+ SD) transformed percentages of snails found in experi- 
mental quadrants over 65 min of sampling at 5-min intervals. 


—#— cysts 

- -& - empty 2 
—><— sediment 
—Ж_ empty 4 


Number of snails 


D 10 15 20 25 30 Sto 40 45 50 55 60 65 


Time (minutes) 


FIG. 4. Numbers of snails found in each experimental quadrant at each sampling time. 


343 


344 PERSSON ET AL. 


At start of the selection experiment, one snail 
that had been starved previously was placed 
in the center of each petri dish. Thereafter, all 
snails were observed continuously, and notes 
were made on which quadrant they were in 
every five min for 65 min. The experimental pro- 
cedure was first tested with a diatiom, Amphora 
Sp., as a food source to verify effectiveness of 
the method. Snails moved randomly until en- 
countering the diatom quadrant and then re- 
mained until cells were consumed. Quantitative 
data for the $. lachrymosa selection experiment 
were recorded as percentages of snails in each 
quadrant at each time. Analysis of variance was 
conducted on transformed (x = arcsinVx) val- 
ues according to Underwood (1997). 

Preliminary, qualitative grazing studies had 
shown that this snail species fed on S. 
lachrymosa cysts when they were presented 
in petri dishes as the only available particle. 
From this preliminary experiment, fecal pellets 
were collected by holding snails between two 
small, 20 um sieves in running seawater over 
night after at least one hour feeding on cysts. 


RESULTS 


In the preliminary, qualitative grazing experi- 
ment, the majority of cysts appeared to be di- 
gested; the fecal pellets were composed of 
partly digested cysts that were rounded, dis- 
colored, and thin-walled without the outer layer 
of calcite crystals, and material that could be 
identified as parts of cysts (starch grains and 
red accumulation bodies) (Fig. 2). 

In the selection experiment, snails were 
significantely (p = 0.0001) more often located 
in the part of the petri dish with cysts, until all 
cysts were consumed (Figs. 3, 4). During the 
65 min observation period, snails moved ran- 
domly and continuously but, upon encounter- 
ing cysts, they slowed and fed on the cysts 
(Fig. 4). Snails returned to the quadrant and 
fed repeatedly until all cysts were consumed. 
One snail did not move at all during the entire 
experiment and had to be removed from the 
data. Only in this petri dish were cysts not 
eaten at all. 


DISCUSSION 


The eastern mud snail cannot be said to be 
an important grazer on dinoflagellate cysts in 
nature because it inhabits the intertidal zone 
(Bianchi & Levinton, 1981), not a habitat where 


dinoflagellate cysts typically accumulate (Dale, 
1983). Nevertheless, this species does feed 
on microphytobenthic algae and settled phy- 
toplankton, and actively forages for these food 
sources on the sediment surface. The objec- 
tive of this experiment was to determine if cysts 
can be attractive food for foraging deposit feed- 
ers, and thus the mud snail is an appropriate 
test grazer. The experiment showed that snails 
were attracted to and consumed Scrippsiella 
lachrymosa cysts. This is in accordance with 
what is known for land plant seeds, which are 
consumed by animals and can be very impor- 
tant food items (Leck et al., 1989, discuss land 
plant seed ecology), because seeds generally 
contain energy stores necessary for germina- 
tion and early development. It is reasonable 
that the same rules of ecology apply in the 
marine sediment as in the soil on land (Pers- 
son, 2000). Resting stages at the sea floor can 
be, and are subjected to grazing pressure. This 
can be assumed to be different for different 
resting stages and also for different animals 
feeding on the same resting stage. Kremp and 
co-workers (2003) fed Scrippsiella lachrymosa 
cysts (the same strain used here) to three dif- 
ferent species of polychaete deposit feeders. 
They found that the cysts passed through the 
worms without being digested. Thus, it appears 
that the mud snails used in the present study 
have digestive capabilities more effective at 
degrading $. lachrymose cysts than do poly- 
chaetes. 

The resting cyst of the toxic (PSP-producing) 
dinoflagellate Alexandrium fundyense has a 
wall that is very resistant to mechanichal dam- 
age, but is not resistant to acids or digestive 
enzymes of many animals. We observed that 
A. fundyense cysts could not be crushed by 
pressing them between two slides (which eas- 
ily crushes other cyst species), but these cysts 
were digested by the eastern oyster, Crasso- 
strea virginica (Persson et al., 2006). We have 
performed another experiment in which A. 
fundyense cysts were fed to mud snails in a 
qualitative, petri-dish experiment in the same 
manner as described here for testing digest- 
ibility of S. lachrymosa cysts. Microscope ob- 
servations of fecal pellets revealed that A. 
fundyense cysts were destroyed in the diges- 
tive process of mud snails; the mucus layer dis- 
appeared, and empty cysts and cysts broken 
at one end were common in fecal pellets. 

We conclude that dinoflagellate cysts can be 
recognized as food by foraging, deposit-feed- 
ing animals and used as a food item. This adds 
to the developing picture of the “seed ecol- 


SNAILS EAT DINOFLAGELLATE CYSTS 345 


ogy” of dinoflagellates. Some species have 
thick-walled cysts and are not easily de- 
stroyed, for example Lingulodinium polyedrum 
(Persson & Rosenberg, 2003); whereas, other 
species are found in higher proportions in an- 
oxic areas, for example, Alexandrium spp. 
(Lewis et al., 1979; Keafer et al., 1992). It is 
logical to conclude that dinoflagellate cysts in 
aerobic areas are subjected to more grazing 
pressure than those in hypoxic or anaerobic 
areas with fewer grazers. Thus, there is a risk 
that cysts of harmful dinoflagellates can accu- 
mulate more if there are more anoxic areas 
with lower grazing pressure on the sea floor. 
Consequently, hypoxia associated with coastal 
eutrophication could be contributing to in- 
creased incidence and severity of harmful al- 
gal blooms by releasing cysts from grazing 
pressure in anoxic benthic habitats. 


ACKNOWLWDGEMENTS 


We are very grateful to Anke Kremp (at 
Woods Hole Oceanographic Institution in 
2002) for providing the Scrippsiella lachrymosa 
B10 strain, along with culturing and encyst- 
ment advice, to Jennifer Alix at Milford Labo- 
ratory for assistance and maintaining the 
culture, and to Hanna and Gunnar Persson 
for collecting snails for the experiment. This 
research was performed while Dr. Agneta 
Persson held a National Research Council 
Research Associateship Award at the National 
Oceanic and Atmospheric Administration, Na- 
tional Marine Fisheries Service Laboratory in 
Milford, Connecticut. Mention of trade names 
does not imply endorsement. 


LITERATURE CITED 


BIANCHI, T. S. & J. S. LEVINTON, 1981, Nutri- 
tion and food limitation of deposit-feeders. Il. 
Differential effects of Hydrobia totteni and 
llyanassa obsoleta on the microbial community. 
Journal of Marine Research, 39 (3): 547-556. 


DALE, B., 1983, Dinoflagellate resting cysts: 
“benthic plankton”. Pp. 69-136, in: G. A. 
FRYXELL, ed., Survival strategies of the algae. 
Cambridge, U.K., Cambridge University Press. 
x + 144 pp. 

GUILLARD R. R. |. & Н. RYTHER, 1962, Stud- 
ies of marine planktonic diatoms. |. Cyclotella 
nana Hustedt and Detonula confervacea 
(Cleve) Gran. Canadian Journal of Microbiol- 
ogy, 8: 229-239. 

KEAFER, В. A., К. O. BUESSELER & D. М. 
ANDERSON, 1992, Burial of living dinoflagel- 
late cysts in estuarine and nearshore sedi- 
ments. Marine Micropaleontology, 20(2): 
147-161. 

KREMP, A., D. H. SHULL & D. M. ANDERSON, 
2003, Effects of deposit-feeder gut passage 
and fecal pellet encapsulation on germination 
of dinoflagellate resting cysts. Marine Ecology 
Progress Series, 263: 65-73. 

LECK, M.A., V. Т. PARKER & К. L. SIMPSON, 
1989, Ecology of soil seed banks. San Diego, 
California, Academic Press. xxii + 462 pp. 

LEWIS, C. M., C. M. YENTSCH:& В. DALE, 1979, 
Distribution of Gonyaulax excavata resting 
cysts in the sediments of Gulf of Maine. Pp. 
235-238, in: Е. J. В. TAYLOR, ed., The biology of 
dinoflagellates. Oxford, U.K., Blackwell Scien- 
tific. xxii + 785 pp. 

PERSSON, A., 2000, Possible predation of cysts 
— a gap in the knowledge of dinoflagellate 
ecology? Journal of Plankton Resarch, 22(4): 
803-809. 

PERSSON, A. 4 R. ROSENBERG, 2003, Impact 
of grazing and bioturbation of marine benthic 
deposit feeders on dinoflagellate cysts. Harm- 
ful Algae, 2(1): 43-50. 

PERSSON, A., В. С. SMITH, С. WIKFORS 8 M. 
QUILLIAM, 2006, Grazing on toxic Alexandrium 
fundyense resting cysts and vegetative cells 
by the Eastern Oyster (Crassostrea virginica). 
Harmful Algae, 5(6): 678-684. 

REID, С.К. 8 С. T. BOALCH, 1987, A new method 
for the identification of dinoflagellate cysts. Jour- 
nal of Plankton Research, 9(1): 249-253. 

SMITH, B. 8 A. PERSSON, 2004, Dinoflagellate 
cyst production in one-liter containers. Jour- 
nal of Applied Phycology, 16(5): 401-405. 

UNDERWOOD, А. J., 1997, Experiments in ecol- 
ogy: their logical design and interpretation us- 
ing analysis of variance. Cambridge, U.K., 
Cambridge University Press. xviii + 504 pp. 


Revised ms. accepted 15 March 2007 


ar a ñas ‘oi 


Er Be RR Ki me ra ee Bose и mul 
ON HUE ue ии err fe Patti 
a к NEE Wa Nei) Sep SENS gi = CNT Lan Ps ee a atic yt sl у AR 

pel a Ne { El me UT M: м Er HE tly 


a. 


p = == 
Tu м ARD ona 
. | ae РЕ A pe Dinas. wit eee TA 
| % р A CA: 
3! | # « + 2 44 Papi bee 
al EL + En $94 an ME 
> ic , 


Pi sens 

t Main EG Dé en 
jas. Speke Ч LT | 

ve SOME: AA af 
LES spa ар Mail In 

La | | pad 4 NAS mi 

he ESTER = Bee ‘<i wey 


Er и An. o A F 
à + Я Maa, * hi Г m В [4 Fe, El on ы : НЯ Or a | { 
à к-р a > a Lt + . и j ae 
ea N Ba De à 4 | м tr) 3 | | р. 
es me i. Er и $ 
E 2 4 Dr, re: SAR dde à 
u + » o > - u 


pak Aka mare ie + és ны аа ан | y 
as site Dies ra e # 

ae а ¡A 
E te BE En sow oe ae Pat | 


инет 3 


a) si ro (Or ent Ame Сок 
АЕ | 
arte El Fr HA ze 


ve gr y 
me 


due as 


1, ex u pd 


г 45 wy 


ee: et Bas oe | 
farm terrine e ter ne 
SE AS RS) oh CS DE 
ani QE alate haber Airing: Pia den р 
- баны ара er PESA U: 
„ ib Fee dienst : р 
гу Acs Be er pres = ¡5 a a éteint Een eg ke hice lees > Li 
La Eee 7 iat Ys. LS Ken IE ee aula als Waly 2! 
ado TL A Chats, EN dr anime x Er ee ir нак sr 
ahs Sr pa a ve aM а ARAS MÍ, ste 


i 
fi 
L 
a | 
is 
à 


Pot 3 


te it ge E 7 DRE fat ER "4 AAA e 
LU A ; , Г 
| | НН 
Е. LT : mn — N > A qu + к | 
| rs | FA ne ee NET eye ad by Eve waste 
eS) а 
RR baat ee Jago pes so cath Ku а sat LEA 


ae ae par : ti | =. k ÿ a Med, NT ER RN р Бы 2 Zu & 
Oral RSC de: a dopa Marie: oa nr que dr à beet ee! À 
ie alé Mut ri ae a sai tet AAA tigen, A sab ' 
ara vale via a Peo cist En Г. aie ciel a 48 0 Lea je PE pio BEA А 
ег Wi a" “i А ‘ish ine vérin «xt er: 7 yin inhi: rene teat a! 
я iii OS Pr ehe nen 
| БД. Lafuente. Rar i, AE A ny! À die 
es ант Peres que E de mies ть beets esi 
ь д. | “y A iy a ee. 


B 
Ei 


= ta LE ps » | Zu 
EST i the ae li E 


in eh macacos hier Gadel Ach on 
mar Het Or GINGEN УВ 
une a Pie trie te rends 
I eee AL ct a habia er я 


MALACOLOGIA, 2008, 50(1-2): 347-350 


ANEW RECORD OF TAE PRESENCE OF 
TWO SPERMATOPHORIC COMPLEXES IN A MALE SHORT-FIN SQUID 
(ILLEX ARGENTINUS, CASTELLANOS, 1960) 


Augusto C. Crespi-Abril 


Centro Nacional Patagonico, Consejo Nacional de Investigaciones Cientificas y Técnicas, 
Boulevard Brown 2825, Puerto Madryn (U9120ACF), Chubut, Argentina; 
crespi@cenpat.edu.ar 


INTRODUCTION 


Several authors have reported morphologi- 
cal abnormalities in cephalopods resulting ei- 
ther from extreme experimental conditions 
(e.g., eggs of cephalopods incubated at tem- 
peratures outside the optimal range for a par- 
ticular species, with increased levels of 
mortality and deformity) (O’Dor et al., 1982; 
Sakurai et al., 1996; Oosthuizen et al., 2002), 
or from unknown causes in nature (i.e., devel- 
opmental abnormalities in embryos and adults 
collected in the field) (Voss, 1957; Bradbury & 
Aldrich, 1971; Gowland et al., 2002). Even 
though numerous malformations have been 
reported, few were localized in the reproduc- 
tive system of cephalopods. For example, 
Hoving et al. (2006) mentioned the presence 
of nidamental glands in males of Ancistro- 
cheirus lesueurii (d’Orbigny, 1842), and Ortiz 
& Ré (2006) described the first case of 
pseudohermaphroditism in Enteroctopus 
megalocyathus (Gould, 1852). To our knowl- 
edge, only one case similar to the one de- 
scribed here has so far been reported, from a 
different species (Bello, 1993). 


RESULTS 


During examination of 2010 Шех argentinus 
males sampled in the San Matias Gulf, Argen- 
tina (40°48’°-42°12’S, 63°44’-65°07’W), from 
April 2005 to August 2007, an anomalous in- 
dividual was found with two spermatophoric 
complexes (Fig. 2): the typical spermatophoric 
complex located on the left side and an atypi- 
cal one on the right side. Both spermatophoric 
complexes showed signs of normal physiologi- 
cal activity, with spermatophores in the sper- 
matophoric (Needham's) sac. The dimensions 
of both spermatophoric complexes were simi- 
lar, except for the length ofthe sperm duct (SD) 


(Table 1). The weight of the typical (10.57 g) 
and atypical (5.4 g) spermatophoric complexes 
were respectively higher and lower than the 
mean weight estimated for this organ from in- 
dividuals of similar maturity condition (95% 
interval of confidence = 7.45—9.97; df = 58). The 
specimen was of 23 cm in mantle length and 
was caught in July 2007 at a depth of 120 m. 
The whole reproductive system of the speci- 
men has been deposited in the marine inver- 
tebrate collection of the Commercial Fish and 
Shellfish Laboratory of the National Patagonian 
Center (LAPEMAR-CENPAT) and is available 
for further examination. 


FIG. 1. Schematic presentation of the macro- 
scopic structure of the spermatophoric complex 
of ommastrephids. SS: spermatophoric sac; SD: 
sperm duct; SG: spermatophoric glands; SpD: 
spermatophoric duct; Te: testis. 


348 CRESPI-ABRIL 


FIG. 2. Internal organs of the abnormal individual (above) showing two spermatophoric 
complexes and of the normal individual (below) showing one spermatophoric complex. 
DG: digestive gland; G: gills; SS: spermatophoric sac; S: stomach; SD: sperm duct; SG: 
spermatophoric glands; Te: testis. 


DOUBLE SPERMATOPHORIC COMPLEX IN /LLEX ARGENTINUS 349 


TABLE 1. Weight and linear measurements of 
the typical (left side of the individual) and the 
atypical (right side of the individual) sperma- 
tophoric complexes (SCo). SS: spermatophoric 
sac; SD: sperm duct; SG: spermatophoric glands; 
SpD: spermatophoric duct. 


Weigth SS SD: 96. Sop 
SCO. (9): (Em): fem => (em): (em) 


Typical 10876010 062210222771 9.98 
Atypical 5.40 65.75 64-260 697 


DISCUSSION 


One aspect of reproduction that all cephalo- 
pods seem to have in common is that they are 
gonochoristic; that is that sex is determined 
genetically; individuals develop as males or 
females and remain the same sex throughout 
their life (Nesis, 1987). In Coleoidea, the re- 
productive system of males is generally formed 
by one duct and one testis. The ectodermal 
part of the duct gives rise to the sperma- 
tophoric complex (SCo) in males and to the 
oviductal glands in females, and the mesoder- 
mal part develops into the proximal vas defer- 
ens in males and proximal oviduct in females 
(Mangold, 1987). The sperm is produced in 
the testis, and mature spermatozoa are shed 
into the proximal portion of the vas deferens 
where, in the convoluted and glandular tubing 
of the spermatophoric organ (sperm duct), they 
are packaged into spermatophores and stored 
in the spermatophoric (Needham’s) sac (Fig. 
1). The terminal portion of the vas deferens 
becomes free from the body surface shortly 
before its ending just within the mantle open- 
ing on the left side (Boyle & Rodhouse, 2005). 
In the particular case of Oegopsida, the male’s 
genital ducts are mostly single and situated on 
the left side, except for members of the family 
Histioteuthidae and the genera Lycoteuthis, 
Selenoteuthis, and Oregoniateuthis of the 
Lycoteuthidae, which possess paired genital 
ducts (Nesis, 1987). Particularly in Calliteuthis 
dofleini (Histoteuthidae), double hectocotyli are 
also present (Nesis, 1987). As mentioned by 
Chun (1910), this condition may perhaps be 
regarded as primary, from which the asymme- 
try of other Oegopsida developed secondarily. 

Amongst the factors affecting the normal 
development of the reproductive system of 
aquatic species, temperature fluctuation 


(Devlin & Nagahama, 2002) and infection by 
viruses and parasites (Pinn et al., 2001) are 
the most thoroughly studied. Particularly in 
mollusks, pollutants are well known for caus- 
ing malformations in the reproductive system 
(i.e., imposex caused by Tributyltin), but their 
effects have been described in detail mainly 
in gastropods (Bryan et al., 1986; Bryan et al., 
1988; Bryan & Gibbs, 1991; Bettin et al., 1996; 
Oberdörster & Cheek, 2001; Goldberg et al., 
2004; Bigatti & Carranza, 2007). There is evi- 
dence that cephalopods can bio-accumulate 
such contaminants as heavy metals from both 
food and water (Miramand & Guary, 1980: 
Miramand & Bentley, 1992). Gerpe et al. 
(2000) have shown that /. argentinus is not an 
exception to this rule; they are bio-accumulat- 
ing heavy metals — Zn, Cu, Cd, and Hg — in 
the digestive gland, gonad and mantle. How- 
ever, for the specimen analyzed in the present 
study, lack of knowledge of possible sources 
of contamination during handling and preser- 
vation would make determinations of pollut- 
ant concentration unreliable. 


ACKNOWLEDGEMENTS 


| am grateful to Paula Sgarlatta and Marlene 
Dherete for their help in the samplings; and to 
Pedro J. Baron and Enrique M. Morsan for the 
comments on the manuscript. 


LITERATURE CITED 


BELLO, G., 1993, Male specimen of Illex coindetii 
(Oegopsida: Ommastrephidae) with two func- 
tional gonoducts. Journal of Cephalopod Biol- 
ogy, 2(2): Notes 1-2. 

BETTIN, C., J. OEHLMANN & E. STROBEN, 
1996, TBT-induced imposex in marine neogas- 
tropods is mediated by an increasing andro- 
gen level. Helgoländer Meeresuntersuchungen, 
50: 299-317. 

BIGATTI, G & A. CARRANZA, 2007, Phenotypic 
variability associated with the occurrence of 
imposex in Odontocymbiola magellanica 
(Gmelin 1791) from Golfo Nuevo. Patagonia. 
Journal of the Marine Biological Association of 
the United Kingdom, 87: 755-759. 

BOYLE, P. & P. RODHOUSE, 2005, Cephalo- 
pods: ecology and fisheries. Abingdon, Oxford, 
Blackwell Publishing. 452 pp. 

BRADBURY, Н. Е. & F. A. ALDRICH, 1971, The 
occurrence of morphological abnormalities in the 
oegopsid squid Шех ilecebrosus (Lesueur, 1821). 
Canadian Journal of Zoology, 49: 377-379. 


350 CRESPI-ABRIL 


BRYAN, G. W. & P. E. GIBBS, 1991, Impact of 
low concentrations of tributyltin (TBT) on ma- 
rine organisms: a review. Pp. 323-362, in: M. 
C. NEWMAN & A. W. MCINTOSH, eds., Metal 
ecotoxicology: concepts and applications. Boca 
Raton, Florida, Lewis Publishers. xvi + 399 pp. 

BRYAN, С. W., Р.Е. GIBBS & С. R. BURT, 1988, 
A comparison of the effectiveness of tri- n- 
butyltin chloride and five other organotin com- 
pounds in promoting the development of 
imposex in the dogwhelk, Nucella lapillus. Jour- 
nal of the Marine Biological Association of the 
United Kingdom, 68: 733-744. 

BRYAN, С. W., Р.Е. GIBBS, |. С. HUMMER- 
STONE & G. R. BURT, 1986, The decline of 
the gastropod Nucella lapillus around south- 
west England: evidence for the effect of 
tributyltin from antifouling paints. Journal of the 
Marine Biological Association of the United 
Kingdom, 66: 611-640. 

CHUN, С., 1910, The Cephalopoda. Part I: 
Oegopsida. 1975 English translation: Jerusa- 
lem, Israel, Keterpress, 2 vols. 436 pp. 

DEVLIN, R. H. & Y. NAGAHAMA, 2002, Sex de- 
termination and sex differentiation in fish: an 
overview of genetic, physiological, and envi- 
ronmental influences. Aquaculture, 208: 191- 
364. 

GERPE, М. S. 3, Е А. ВЕ MORENO, \ J: 
MORENO & М. L. PATAT, 2000, Cadmium, zinc 
and copper accumulation in the squid //lex 
argentinus from the southwest Atlantic coast. 
Marine Biology, 136: 1039-1044. у 

GOLDBERG, R., А. AVERBUJ, М. CLEDON, D. 
LUZZATTO & N. SBARBATI NUDELMAN, 
2004, Search for triorganotins along the Mar 
del Plata (Argentina) marine coast: finding of 
tributyltin in egg capsules of a snail Adelomelon 
brasiliana (Lamarck 1822) population showing 
imposex effects. Applied Organometallic 
Chemistry, 18: 117-123. 

GOWLAND, Е. С., М. A. MOLTSCHANIWSKYJ 
& M.A. STEER, 2002, Description and quanti- 
fication of developmental abnormalities in a 
natural Sepioteuthis australis spawning popu- 
lation (Mollusca: Cephalopoda). Marine Ecol- 
ogy Progress Series, 243: 133-141. 

НОММС, H.. JT, М. А.С. ROELEVELD, М. В. 
LIPINSKI & J. J. VIDELER, 2006, Nidamental 
glands in males of the oceanic squid Ancistro- 
cheirus lesueurii (Cephalopoda: Ancistrocheiri- 
dae) — sex change or intersexulity? Journal of 
Zoology, 269(3): 341-348. 


MANGOLD, K., 1987, Reproduction. Pp. 157- 
200, in: P.R. BOYLE, ed., Cephalopod life cycles. 
Volume Il: comparative reviews. London, Aca- 
demic Press. xxi + 441 pp. 

MIRAMAND, P. & D. BENTLEY, 1992, Concen- 
tration and distribution of heavy metals in tis- 
sues of two cephalopods, Eledone cirrhosa and 
Sepia officinalis, from the French coast of the 
English Channel. Marine Biology, 114: 407- 
414. 

MIRAMAND, P. & J. С. GUARY, 1980, High con- 
centrations of some heavy metals in tissues of 
the Mediterranean octopus. Bulletin of Envi- 
ronmental Contaminants and Toxicology, 24: 
183-788 

NESIS, K. N., 1987, Cephalopods of the world: 
squids, cuttlefishes, octoposes, and allies. 
Neptune City, New Jersey, T. F. H. Publications. 
350 pp. [translated from Russian]. 

OBERDORSTER, Е. &A. O. CHEEK, 2001, Gen- 
der benders at the beach: endocrine disruption 
in marine and estuarine organisms. Environ- 
mental Toxicology and Chemistry, 20: 23-36. 

O'DOR, R. K., N. BALCH, Е. А. FOY, В. W. M. 
HIRTLE, О.А. JOHNSTON & T. AMARATUNGA, 
1982, Embryonic development of the squid, Illex 
illecebrosus, and effect of temperature on de- 
velopment rates. Journal of Northwest Atlantic 
Fishery Science, 3: 41-45. 

OOSTHUIZEN, A., M. ROBERTS & W. SAUER, 
2002, Temperature effects on the embryonic 
development and hatching success of the squid 
Loligo vulgaris reynaudii. Bulletin of Marine 
Science, 71(2): 619-632. 

ORTIZ, N. & M. E. RE, 2006, First report of 
pseudohermaphroditism in cephalopods. Jour- 
nal of Molluscan Studies, 72: 321-323. 

PINN, Е. H., В. J. A. ATKINSON & A. ROGER- 
SON, 2001, Sexual dimorphism and intersexu- 
ality in Upogebia stellata (Crustacea: Decapoda 
Thalassinidae). Journal cf the Marine Biologi- 
cal Association of the United Kingdom, 81: 
1061-1062. 

SAKURAI, Y., J. R. BOWER, Y. AKAMURA, S. 
YAMAMOTO & K. WATANABE, 1996, Effect of 
temperature on development and survival of 
Todarodes pacificus embryos and paralarvae. 
American Malacological Bulletin, 13(1/2): 89- 
95 


VOSS, С. L., 1957, Observation on abnormal 
growth of the arms and tentacle in the squid 
genus Rossia. Quartertly Journal of the Florida 
Academy of Sciences, 20: 129-132. 


Revised ms. accepted 22 February 2008 


MALACOLOGIA, 2008, 50(1-2): 351-357 


LACK OF MITOCHONDRIAL DNA DIVERSITY IN INVASIVE 
APPLE SNAILS (AMPULLARIIDAE) IN HAWAII 


Chuong T. Tran!” Kenneth A. Hayes'* & Robert H. Cowie™ 


ABSTRACT 


Three species of apple snails (Ampullariidae) have been introduced to Hawaii. In order 
to clarify their identities, determine their geographic origins, and evaluate their mtDNA 
diversity in Hawaii, we sequenced the COI and ND6 mtDNA markers from 103 snails col- 
lected on the six main Hawaiian islands. Our samples included Pila conica and Pomacea 
canaliculata, whose identities were confirmed by phylogenetic analysis that included other 
ampullariid species. The third species, Pomacea diffusa, known to have been present in 
the past, was not found. Neither species exhibited any variation at either marker. Both 
species may have been introduced to Hawaii as single introductions, possibly from the 


Philippines. 


Key words: Pomacea, Pila, mtDNA, invasive species. 


INTRODUCTION 


Invasive species are a major concern be- 
cause of their potential for great ecological, 
agricultural, human health, and economic im- 
pacts (Mack et al., 2000). These alien species 
may affect native species through predation, 
competition, introduction of disease, and hy- 
bridization (OTA, 1993). Well-known examples 
include the zebra mussel (Dreissena poly- 
morpha), introduced to North America and one 
of the main factors endangering native fresh- 
water unionid clams, and predatory snails 
(Euglandina rosea and others) introduced to 
Pacific and Indian Ocean islands, where they 
have caused the decline and extinction of na- 
tive snails (Lydeard et al., 2004). 

The Ampullariidae include a number of com- 
plexes of more or less cryptic species. There 
are nine recognized genera, the largest of 
which, Pomacea Perry, 1810, has 117 nomen- 
claturally valid species (Cowie & Thiengo, 
2003). The real number of species may be 
closer to 50, as many synonyms have prob- 
ably been generated based on minor varia- 
tions in shell characteristics that may be 
phenotypically plastic (Estebenet & Martin, 
2003). These freshwater snails are known as 
“apple snails” because of the large, round and 
sometimes greenish appearance of the shells 


in some genera, including Pomacea (Cowie & 
Thiengo, 2003). Their biology, including ecol- 
ogy, life-history and behavior, have been re- 
viewed by Cowie (2002). 

Native South American apple snails in the 
genus Pomacea were introduced to Taiwan in 
1980 with the intention of developing them as 
a human food resource, both locally and for 
the international gourmet trade (Mochida, 
1991). They were later introduced to other parts 
of Asia, New Guinea, Guam, and Hawaii, but 
their development as a food resource was only 
partially successful, and the snails either es- 
caped or were released and have become 
major agricultural pests, notably in rice and taro, 
as well as other crops (Mochida, 1991; Cowie, 
2002; Lai et al., 2005; Joshi & Sebastian, 2006). 
Some species are also common in the aquar- 
ium trade, including in Hawaii. In Hawaii, apple 
snails are now found in the wild predominantly, 
but not exclusively, in taro-growing areas 
(Cowie, 1995, 1996; Cowie et al., 2007), where 
they cause major crop damage (Levin et al., 
2006). The environmental consequences of the 
snails becoming established in Hawaii have not 
been evaluated, but their voracious and gen- 
eralist feeding habits (Lach et al., 2001), com- 
bined with past experience with other 
introduced species in the Hawaiian Islands 
(Staples & Cowie, 2001) and their known eco- 


‘Center for Conservation Research and Training, Pacific Biosciences Research Center, University of Hawaii, 3050 Maile Way, 


Gilmore 408, Honolulu, Hawaii 96822, U.S.A. 


“Biology Program, University of Hawaii, Honolulu, Hawaii 96822, U.S.A. 
¿Department of Zoology, University of Hawaii, Honolulu, Hawaii 96822, U.S.A. 


“Corresponding author: cowie@hawaii.edu 


332 TRAN ETAL. 


(4/2) 


2) (412) 
A (412) 


«му Kauai 


Oahu 


(10/3) (4/3) m» 
L 


(211) 


50 km 


{4} 
DS gy 
{4} (272) 
ES | / 
(2} 
De ie 
= 


ФЕ 


anai 
(4/2) es (8/1) 


(311) 


Hawail 


FIG. 1. The Hawaiian Islands showing locations of sampling sites of the 2004-2005 survey from 
which snails were sequenced (Pila conica only on Molokai; Pomacea canaliculata on all islands 
except Molokai). Numbers in parentheses are the number of COI sequences followed by the number 
of ND6 sequences from each site; single numbers are from sites from which only COI was sequenced; 
there were no sites from which ND6 only was sequenced. 


logical impacts elsewhere (Carlsson et al., 
2004), suggest that major damage, including 
destruction of native vegetation and competi- 
tion with native freshwater fauna is possible. 

Because of their potential negative impacts, 
it is essential to identify introduced species 
reliably (Holland et al., 2004). Morphological 
characteristics are often adequate for species 
identification, but molecular data are neces- 
sary to identify cryptic species that cannot be 
distinguished using morphological character- 
istics alone. This is especially the case for 
Pomacea, because the overall highly con- 
served external morphology across the genus 
yet considerable intraspecific shell variation 
obscures the true number of species and their 
identities (Cazzaniga, 2002). In addition, 
knowledge of their geographic origins and of 
their phylogeography in the introduced envi- 
ronment are also potentially important in un- 
derstanding and managing these invasive 
snails. Molecular data can more readily ad- 
dress such questions. By sequencing the cyto- 
chrome с oxidase subunit | (СО!) and NADH 
dehydrogenase subunit 6 (ND6) mitochondrial 


DNA markers this study aimed to confirm the 
identities of the introduced ampullariids in 
Hawaii, determine their probable origin, and 
investigate any geographic genetic structur- 
ing of the introduced populations. 


MATERIALS AND METHODS 


As part of a survey of apple snail distribution 
in the Hawaiian Islands during 2004-2005 
(Cowie et al., 2007) apple snails were collected 
from 21 sites on six of the main Hawaiian Is- 
lands. Details of these sites and collections are 
given by Cowie et al. (2007). Five additional 
collections were obtained from pet stores, an 
Asian food market in Honolulu, and from other 
collectors who gave Honolulu as the locality 
(Fig. 1). All snails were processed and total 
DNA extracted following Hayes et al. (in press). 
Amplification of a 658 bp portion of the mito- 
chondrial DNA (mtDNA) encoded cytochrome 
с oxidase subunit | (COI) gene and a 607 bp 
fragment of the NADH dehydrogenase subunit 
6 (ND6) gene was carried out following stan- 


LACK OF DNA DIVERSITY IN INVASIVE APPLE SNAILS 399 


Cipangopaludina chinensis 


au Pila polita 
| 100/98 z 
99/96 № Pila sp. 
99/96 = A 
Pila conica 
aa Pomacea paludosa 
— P. insularum E 
Clade A 
71/88 
0.02 | warm 
400 
Y 06/66 50 
1007100 | eee 
TOUT Jos Р canaliculata 
| | por Hawaii 
ER CladeB 
| | Argentina 
100/99 ms 
_ 10079 р dolioides 
Other Southern and 
| 100/87 Р haustrum Eastem Asia Countries 
| /86 007100 P. diffusa 
Eu ИИ Р bridgesii 
73/61 Р scalaris 
Marisa comuanelis 


FIG. 2. ACOI mtDNA minimum evolution tree and a haplotype network (95% parsimony limit). The 
phylogenetic tree illustrates the relationships of the two Hawaiian apple snail species, Pomacea 
canaliculata (Clades A and B) and Pila conica, to other ampullariid taxa. Terminal branches for all 
species, except P. canaliculata, have been collapsed in order to simplify the tree presentation. Node 
values represent 500 bootstrap replicates (> 50%) under ME/MP. The haplotype network illustrates 
the genealogical relationships among P. canaliculata individuals in Clade A, the only clade containing 
Hawaiian samples. The Clade B network (not shown) is composed of haplotypes found in the Philip- 
pines, Argentina and a number other Asian countries, the closest one of which is an additional 22 


steps from the closest haplotype in Clade A. 


dard polymerase chain reaction protocols and 
primers from Folmer et al. (1994) and Hayes 
(in prep). Cycle-sequencing, electrophoresis 
and analyses were carried out on an ABI 
3730XL, and all unique Hawaiian sequences 
generated in this study have been deposited 
in Genbank (Accession nos. EU523129- 
EU523131). To definitively identify the Hawai- 
ian snails, all COl sequences were combined 
with a subset of COI sequences from south- 
ern and eastern Asia, the U.S.A., and South 
America (Hayes et al., in press), and phyloge- 
netic analysis was carried out under minimum 
evolution (ME) and maximum parsimony (MP) 
search criteria in PAUP (Swofford, 2003) with 


500 bootstrap replicates. Distances for mini- 
mum evolution searches were determined us- 
ing the best fit maximum likelihood model of 
substitution in MODELTEST (Posada & 
Crandall, 1998). The ND6 sequences were also 
compared in a phylogenetic framework to one 
another and to other samples from the Philip- 
pines and the native range in Argentina. Also, 
COI haplotypes were used to construct a par- 
simony network with a 95% parsimony limit in 
TCS in order to illustrate the relationships 
among the Hawaiian samples and those from 
other regions. The lack of variation within apple 
Snail species in Hawaii (See Results) prevented 
further population level analyses. 


354 TRAN ETAL. 


RESULTS 


Based on their morphology and by reference 
to previous published records of ampullariids 
in Hawaii (Cowie, 1995, 1996, 1997; Cowie et 
al., 2007), two species were found: Pila conica 
(Gray, 1828), which is native to south-east 
Asia, and Pomacea canaliculata (Lamarck, 
1822), native to South America. The former 
was found only on Molokai, the latter on all 
islands except Molokai. The specific identity 
of the latter was confirmed by phylogentic com- 
parison with СО! sequences from material 
collected from other locations, as stated above 
(Fig. 2; Rawlings et al., 2007; Hayes et al., in 
press). A third species, Pomacea diffusa 
(Blume, 1957), previously known from Hawaii 
although incorrectly identified as Pomacea 
bridgesii (Reeve, 1856), was not found. 

We sequenced COI from 14 Pila conica and 
89 Pomacea canaliculata, representing every 
site, and ND6 from 33 P. canaliculata, repre- 
senting 14 of the 21 sites at which it was found 
(Fig. 1). All individuals of Pomacea canaliculata 
shared a single COI haplotype and all Pila 
conica also shared a single (different) COI 
haplotype. The Pomacea canaliculata also all 
exhibited only a single ND6 haplotype, also 
shared with samples from the Philippines. All 
analyses yielded congruent phylogenetic trees 
with only minor differences in support values 
for deeper nodes (Fig. 2). 


DISCUSSION 


Pomacea canaliculata was first recorded in 
Hawaii in 1989, from the island of Maui and 
subsequently on all main islands except 
Molokai (Cowie, 1995, 1996), although it may 
have been present before this (Levin et al., 
2007). These snails are popular as a food item 
among the Filipino community in Hawaii, and 
there are consistent and extensive anecdotal 
reports that the Philippines were the source 
of the introductions to Hawaii (Levin et al., 
2007). The single СО! haplotype shared by 
all 89 Pomacea canaliculata is also shared by 
some individuals from Argentina, part of its 
native range, and is the most common haplo- 
type for this snail in the Philippines (Fig. 2; 
Hayes et al., in press). The same is true for 
ND6. Our results are therefore consistent with 
the explanation that P. canaliculata was 
brought to Hawaii from the Philippines, and 


for the same reason it was initially taken to 
Asia, as a human food resource. The Р. 
canaliculata from pet stores in Hawaii also 
share the same COI haplotype and therefore 
probably originated locally from one of the pre- 
viously introduced populations. Although this 
is the most common and widespread haplo- 
type in the Philippines, there are numerous 
other haplotypes there. The presence of only 
this one haplotype in Hawaii therefore sug- 
gests a single introduction. 

Pila conica, was first recorded also from 
Maui, in 1966, and in 1991 from Oahu and 
Molokai (Cowie, 1995). During our survey we 
only found it on Molokai (Cowie et al., 2007), 
from which Р canaliculata is absent. All 14 
snails sequenced possessed identical COI 
haplotypes, indicating that the populations on 
Molokai may have resulted from a single in- 
troduction. This species is also known from 
the Philippines, where it is native, and it may 
have been introduced by the same route and 
for the same purpose as Pomacea canalicula- 
fa. 

An alternative explanation for the lack of 
mitochondrial genetic variation in the intro- 
duced Hawaiian populations could be that 
strong selection for a single haplotype has led 
to removal of all others from the populations, 
a selective sweep. This scenario seems highly 
unlikely, since it would have had to have taken 
place on multiple islands and in multiple sepa- 
rate populations within islands, and in two spe- 
cies. 

Since ND6 is a faster-evolving gene than 
COI, particularly in snails (Satler & Steiner, 
2004), its lack of variation further supports the 
interpretation that all individuals of each of the 
two species are very closely related and that 
each species may have been introduced just 
once and from a single population. 

The lack of mitochondrial diversity in these 
introduced populations is surprising, given that 
P. canaliculata especially is an extremely suc- 
cessful invader in Hawaii, and the fact that 
multiple introductions have occurred in south- 
ern and eastern Asia (Hayes et al., in press). 
Colonization by alien species has often been 
thought to involve loss of allelic diversity re- 
sulting from a genetic bottleneck because of 
the low numbers of colonists, which in turn 
may lead to a further reduction in genetic di- 
versity as a result of increased inbreeding. We 
recorded no diversity at two mtDNA loci, and 
in principal such a bottleneck would also af- 


LACK OF DNA DIVERSITY IN INVASIVE APPLE SNAILS 355 


fect nuclear loci, although to a lesser degree. 
Such loss of diversity can lead to two conse- 
quences. First, population growth is limited 
and the likelihood of its survival is low 
(Ellstrand & Elam, 1993). Second, low genetic 
diversity limits evolution in the population 
(Sakai et al., 2001). There is often then a lag 
between initial colonization and proliferation 
and spread, during which evolution leads to 
adaptation to the new habitat, development 
of invasive life-history characters, or purging 
of the genetic load of inbreeding depression; 
the genetic constraints are overcome (Mack 
et al., 2000; Sakai et al., 2001). Although 
Pomacea canaliculata may have been intro- 
duced to Hawaii earlier in the 1980s than the 
first vouchered record indicates, it has not 
exhibited a significant lag time, but spread 
rapidly and became a serious pest well within 
a decade following its introduction (Lach & 
Cowie, 1999). 

Propagule pressure, or the number of sepa- 
rate introductions, is recognized as important 
in the successful establishment of many intro- 
duced species. For example, the European 
starling (Sternus vulgaris) and the house spar- 
row (Passer domesticus) became widely in- 
vasive in North America only after repeated 
introductions (Ehrlich, 1989; Sakai et al., 
2001). Multiple introductions may provide the 
genetic variation needed for many successful 
invasions (Lavergne & Molofsky, 2007; Roman 
& Darling, 2007) and may have contributed to 
the success of Pomacea spp. introduced to 
southern and eastern Asia (Hayes et al., in 
press). Roman & Darling (2007) have argued 
that the paradox of the success of invasive 
species despite low genetic diversity is in fact 
a “paradox lost”, inasmuch as in many cases 
multiple introductions led to no loss of genetic 
diversity, and in those cases exhibiting reduced 
diversity, their success could be explained in 
other ways, including the possibility that mo- 
lecular analyses may underestimate non-neu- 
tral genetic diversity, that which would permit 
success of an invasive species, and that re- 
productive mode may be important: species 
able to reproduce asexually might tolerate loss 
of genetic diversity more readily. This latter 
may be the case in invasive freshwater clams 
in the genus Corbicula, of which a single clonal 
lineage extends from Michigan to Patagonia 
(Lee et al., 2005). 

Some instances of reduced diversity can be 
explained on a case-by-case basis. For in- 


stance, introduced Argentine ants (Linepithema 
humile) exhibit much reduced genetic varia- 
tion, but this is the key to their success as an 
invasive species (Tsutsui et al., 2000), because 
reduced variation permits the formation of su- 
per-colonies that are more successful than 
normal colonies because of the reduction in 
costs associated with territoriality. 

However, while we were not able to assess 
non-neutral genetic diversity, ampullariids are 
obligately sexual reproducing species with 
separate sexes. It indeed remains paradoxi- 
cal, then, that Pomacea canaliculata and to a 
lesser extent Pila conica are such successful 
invaders in Hawaii, despite their extremely re- 
duced mitochondrial variation. Similarly, the 
success of the invasive freshwater snail 
Ferrissia fragilis in Europe, which exhibits little 
genetic variation in its introduced range, re- 
mains unexplained despite speculation that its 
biological attributes may be important (Walther 
et al., 2006). Equally, reasons for the success 
of the introduced Indo-Pacific fish Fistularia 
commersonii in the Mediterranean despite the 
presence of only two mtDNA haplotypes are 
unknown (Golani et al., 2007). 

However, it may not be necessary to invoke 
multiple introductions to account for their suc- 
cess. Preliminary analyses of mitochondrial 
diversity in the native ranges of various 
Pomacea species show that frequent bottle- 
necks have played a role in structuring popu- 
lations (Hayes et al., in press). This may have 
permitted Pomacea canaliculata and other in- 
vasive species to become successful despite 
reduced genetic diversity. 

As Roman & Darling (2007) said, “although 
measurements of relative genetic diversity are 
becoming more common, research into the 
mechanisms explaining the success of low-di- 
versity populations lags behind considerably”, 
contradicting their “paradox lost” assertion. 
Research to further our understanding of how 
genetic diversity is related to invasiveness re- 
mains crucial in development of management 
strategies for alien species. 


ACKNOWLEDGEMENTS 


We thank the numerous people who have 
collected comparative material for us in the past, 
especially Ravi Joshi and Silvana Thiengo. 
Funding was provided by the U.S. Department 
of Agriculture. 


356 TRAN ETAL. 


LITERATURE CITED 


CARLSSON, М. O. L., С. BRONMARK €. L.-A. 
HANSSON, 2004, Invading herbivory: The 
golden apple snail alters ecosystem function- 
ing in Asian wetlands. Ecology, 85: 1575-1580. 

CAZZANIGA, N. J., 2002, Old species and new 
concepts in the taxonomy of Pomacea (Gas- 
tropoda: Ampullariidae). Biocell, 26: 71-81. 

COWIE, R. H., 1995, Identity, Distribution and 
Impacts of Introduced Ampullariidae and 
Viviparidae in the Hawaiian Islands. Journal of 
Medical and Applied Malacology, 5 [issue for 
1993]: 61-67. 

COWIE, R. H., 1996, New records of introduced 
land and freshwater snails in the Hawaiian Is- 
lands. Bishop Museum Occasional Papers, 46: 
25-27. 

COWIE, R. H., 1997, Catalog and bibliography 
of the nonindigenous nonmarine snails and 
slugs of the Hawaiian Islands. Bishop Museum 
Occasional Papers, 50: 1-66. 

COWIE, R. H., 2002, Apple snails (Ampullariidae) 
as agricultural pests: their biology, impacts and 
management. Pp. 145-192, in: G. M. BARKER, 
ed., Molluscs as crop pests. Wallingford, U.K., 
CABI Publishing. xii + 468 pp., [4] pp. of pls. 

COWIE, R. H. & S. C. THIENGO, 2003, The 
apple snails of the Americas (Mollusca: Gas- 
tropoda: Ampullariidae: Asolene, Felipponea, 
Marisa, Pomacea, Pomella): a nomenclatural 
and type catalog. Malacologia, 45: 41-100. 

COWIE, R. H., K. A. HAYES, C. T. TRAN & P. 
LEVIN, 2007, Distribution of the invasive apple 
snail Pomacea canaliculata (Lamarck) in the 
Hawaiian Islands (Gastropoda: Ampullariidae). 
Bishop Museum Occasional Papers, 96: 48-51. 

EHRLICH, P. R., 1989, The attributes of invad- 
ers and the invasion processes: vertebrates. 
Pp. 315-328, in: J. A. DRAKE, H. A. MOONEY, Е. DI 
CASTRI, R. H. GROYAS, F. J. KRUGER, M. REJMÁNEK 
& M. WILLIAMSON, eds., Biological invasions: a 
global perspective. Chichester, New York, John 
Wiley 8 Sons. xxix + 535 pp. 

ELLSTRAND, М. С. 8 D. К. ELAM, 1993, Popu- 
lation genetic consequences of small popula- 
tion size: implication for plant conservation. 
Annual Review of Ecology and Systematics, 
24: 217-242. | 

ESTEBENET, А. L. & Р. В. MARTIN, 2003, Shell 
interpopulation variation and its origin in 
Pomacea canaliculata (Gastropoda: Ampullarii- 
dae) from southern Pampas, Argentina. Jour- 
nal of Molluscan Studies, 69: 301-310. 

FOLMER, O., M. BLACK, W. НОЕН, К. LUTZ & 
R. VRIJENHOEK, 1994, DNA primers for am- 
plification of mitochondrial cytochrome c oxi- 
dase subunit | from diverse metazoan 
invertebrates. Molecular Marine Biology and 
Biotechnology, 3: 291-299. 

GOLANI, D., Е. AZZURRO, M. CORSINI-FOKA, 
M. FALAUTANO, F. ANDALORO & G. BER- 
NARDI, 2007, Genetic bottlenecks and suc- 
cessful biological invasions: the case of a 
recent Lessepsian migrant. Biology Letters, 
published online: doi:10.1098/rsbl.2007.0308 


HAYES, К. A., К. С. JOSHI, S. С. THIENGO €: 
R. H. Cowie, in press, Out of South America: 
multiple origins of non-native apple snails in 
Asia. Diversity and Distributions. 

HOLLAND, B., М. DAWSON, С. CROW & D. 
HOFMANN, 2004, Global phylogeography of 
Cassiopea (Scyphozoa: Rhizostomeae): mo- 
lecular evidence for cryptic species and mul- 
tiple invasions of the Hawaiian Islands. Marine 
Biology, 145: 1119-1128. 

JOSHI, R. C. & L. S. SEBASTIAN, eds., 2006, 
Global advances in ecology and management 
of golden apple snails. Philippine Rice Re- 
search Institute, Nueva Ecija. x + 588 pp. 

LACH, L. & R. H. COWIE, 1999, The spread of 
the introduced freshwater apple snail Pomacea 
canaliculata (Lamarck) (Gastropoda: Ampul- 
lariidae) on O’ahu, Hawai'i. Bishop Museum 
Occasional Papers, 58: 66-71. 

LACH, L., D. К. BRITTON, К. J. RUNDELL € В. 
H. COWIE, 2001, Food preference and repro- 
ductive plasticity in an invasive freshwater 
snail. Biological Invasions, 2(4): 279-288. 

LAI, P.-Y., У. Е. CHANG & R. H. COWIE, eds., 
2005, Proceedings - APEC symposium on the 
management of the golden apple snail, Sep- 
tember 6-11, 2004. National Pingtung Univer- 
sity of Science and Technology, Chinese Taipei. 
[Mi]. 153.0. 

LAVERGNE, S. & J. MOLOFSKY, 2007, In- 
creased genetic variation and evolutionary 
potential drive the success of an invasive grass. 
Proceedings of the National Academy of Sci- 
ences, 104: 3883-3888. 

LEE, Т., $. SIRIPATTRAWAN, С. ITUARTE & D. 
O FOIGHIL, 2005, Invasion of the clonal clams: 
Corbicula lineages in the New World. Ameri- 
can Malacological Bulletin, 20: 113-123. 

BEVIN PR: IRA COWIE J. TAYLOR K. 
BURNETT, K. A. HAYES & C. FERGUSON, 
2006, Apple snail invasions and the slow road 
to control: ecological, economic, agricultural 
and cultural perspectives in Hawaii. Pp. 325- 
335, п: R. С. JOSHI & |. С. SEBASTIAN, eds., Glo- 
bal advances т ecology and management of 
golden apple snails. Philippine Rice Research 
Institute, Mufoz, Nueva Ecija. x + 588 pp. 

LYDEARD, С., К. H. COWIE, W. Е. PONDER, A. 
E. BOGAN, P. BOUCHET, S. CLARK, K. S. 
CUMMINGS, T. J. FREST, О. GARGOMINY, 
DICMERBERIMR HERSHLER, K. PEREZ. 
В. ROTH, M. SEDDON, Е. E. STRONG € F. С. 
THOMPSON, 2004, The global decline of non- 
marine mollusks. BioScience, 54: 321-330. 

MACK, R. N., D. SIMBERLOFF, W. M. LONS- 
DALE, H. EVANS, M. CLOUT & Е. А. BAZZAZ, 
2000, Biotic invasions: causes, epidemiology, 
global consequences, and control. Ecological 
Applications, 10: 689-710. 

MOCHIDA, O., 1991, Spread of freshwater 
Pomacea snails (Pilidae, Mollusca) from Argen- 
tina to Asia. Micronesia, Supplement 3: 51-62. 

OTA [Office of Technology Assessment], 1993, 
Harmful non-indigenous species in the United 
States. Washington, D.C., U.S. Government 
Printing Office. viii + 391 pp. 


LACK OF DNA DIVERSITY IN INVASIVE APPLE SNAILS Sor 


POSADA, D. & K. A. CRANDALL, 1998, Model- 
test: testing the model of DNA substitution. 
Bioinformatics, 14: 817-818. 

RAWLINGS, T.A., К. А. HAYES, К. H. COWIE 8 
T. M. COLLINS, 2007, The identity, distribu- 
tion, and impacts of non-native apple snails in 
the continental United States. BMC Evolution- 
ary Biology, 7: 97 [14 p.] 

ROMAN, J. & J. A. DARLING, 2007, Paradox lost: 
genetic diversity and the success of aquatic 
invasions. Trends in Ecology and Evolution, 22: 
454-464. 

SAKAI, А. K., Е. W. ALLENDORF, J. $. HOLT, D. 
M, LODGE... MOLOFSKY КГА: WITH 5. 
BAUGHMAN, R. J. CABIN, J. E. COHEN, N. 
С. ELLSTRAND, О.Е. MCCAULEY, P. O’NEIL, 
|. М. PARKER, J. М. THOMPSON & $. С. 
WELLER, 2001, The population biology of т- 
vasive species. Annual Review of Ecology and 
Systematics, 32: 305-332. 

SATLER, M. & G. STEINER, 2004, Comparison 
of substituion rates in molluscan mitochondrial 
genes. P. 131, in: Е. Е. WELLS, ed., Molluscan 
megadiversity: sea, land and freshwater. World 
Congress of Malacology, Perth, Western Aus- 


tralia. 11-16 July 2004. Perth, Western Aus- 
tralian Museum. xlvili + 187 pp. 

STAPLES, G. W. & R. H. COWIE, eds., 2001, 
Hawai'i’s invasive species. a guide to invasive 
plants and animals in the Hawaiian Islands. 
Honolulu, Mutual Publishing & Bishop Museum 
Press. xii + 116 p. 

SWOFFORD, D. L., 2003, PAUP*. Phylogenetic 
Analysis Using Parsimony (‘and other meth- 
ods). Version 4. Massachusetts, Sinauer As- 
sociates, Sunderland. 

TSUTSUI, N. D., A. V. SUAREZ, D. A. HOLWAY 
& T. J. CASE, 2000, Reduced genetic varia- 
tion and the success of an invasive species. 
Proceedings of the National Academy of Sci- 
ences, 97: 5948-5953. | 

WALTHER, A. C., T. LEE; J.B. BURCH & D. О 
FOIGHIL, 2006, Confirmation that the North 
American ancylid limpet Ferrissia fragilis 
(Tryon, 1863) is a cryptic invader of European, 
and East Asian, freshwater ecosystems. Jour- 
nal of Molluscan Studies, 72: 318-321. 


Revised ms. accepted 14 February 2008 


| bles et: ré zy RES en ee 
i den pta Pe Apra Men 


= LANA oa de 


| PE A 
He DER 


8 1 
rois 25: re on 
KA arene CT TEAM RP 
a = ti the ee :. 
eut FE acta in Te 
po Prop Al € Len MN Patan oe 
и СН Por: os The sts Stent foe: ts sree 
ere. He toy ve ee tétanie | 
A ETAT, be ee 
| ит ul EL. 2 LR ds, Е. RASH, si RE Лех 
км vale Ario, are. fee RDA à 
р Cines, Nom YORE, У 
Pind F 
PHOS, Pape 


2. к 

I ARAN i. oe Re ELN 

PONT RENNES BE nail NES 

ut a. picado Па 

At ae ie as Gagingy вое. м Ее: rent, 
a RUE ine, 200$ Sha 

Е Pr 0h ad es PU if 


xi rience en EA jalo Dista iz RON ES 
a) Parr осы. re ve У: 


As BIER ов 2 
PONE: fl. 02 Gi doc. A. IH. В oe 
ee 


nl > ae rubs 


и, E Е Hi 


EEK a 


A. NEN, VERA: Ne 
i И noe ae ey 


Ei р u 
une = a _ m E mi 
я ER 
7 h h = o A =. 


on Ca 


pe (REIS SIRCATMANAS € 


a vr ve 
> ВАА Sere 


er ie De 
АС 
ee er 


gu merida and eg 


| ne = aa E 
ga ae Bere nfs 385 de ew Ej ae at = “ig 
Pe MD al NA ny Reihe AA. 
a e ch the Matioge! Aunt of Sch 
Penes, | | 


AA. - 


: CM : Fr 
ire LL ef 


Y Fl ene BUGS, BBC Of EU 


CON AO iles 20 13.127: + aie 
LEWIN PB НСО MES TALA O - 
ASE 43) A: HAVES: SC) FERN ENS 
ОБА SAME Hairs «ht so a. ПВ 
Ba ı ces HOMO. aa i 
AL Leurs ye EN A ger En Le D an 
Se, if À IN à : eases at: AS E 
bar ‘ava “eal y ec e Md dé e a of A 
Pac Zr SEE! д HR ram 2 ОА 
inde: Fer svg CR 
ИОРАН ВК ЗЕ Wi PONDER 4 
Е DERE À А 5. CLARK, ICE 
MAIS БЕСТ + Ci NT 
Die HERBERT R MEREMLER Y TEREZL 
RON M SE BOCH E SINO ENS 
THOMPAG le За р artes” nur TUS y he 
a Acs. Бросок, Gt 501 33 
MALE x. $ СЕНЕ: ЖЕ AE 
| De EVANS LITER GA IZAZ, 
"Y Bir. swaront cn: at À нии к 
: = sae se DUR, Bee) Cortes: Enola 
ia ces. | e У 


Bi 1. 
Espais (PR 


my won. 


senna. SENOS, + 


aire Aalen pe MT CR y Мег. ri 
Funk ROT 
TASTE : + 


Soc) SIMS CS 
ЧАЯ niche, aa in. ire ici Webs. AD oa 


LETTER TO THE:EDITOR 


MALACOLOGIA, 2008, 50(1-2): 361 


PUBLICATION DATES OF MALACOLOGIA VOLUMES AND ISSUES 


Vol./Issue Cover Date Publication Date  Vol./Issue Cover Date Publication Date 
1(1) Oct. 1962 14 Nov. 1962 2202) 1982 24 June 1982 
1(2) July 1963 7 Aug. 1963 23(1) 1982 18 Aug. 1982 
1(3) June 1964 1 June 1964 23(2) 1983 28 Feb. 1983 
2(1) Sept. 1964 22 Sept. 1964 24(1-2) 1983 29 Sept. 1983 
2(2) Feb. 1965 25 Feb. 1965 25(1) 1984 29 March 1984 
2(3) April 1965 29 April 1965 25(2) 1984 29 Aug. 1984 
3(1) Aug. 1965 31 Aug. 1965 26(1-2) 1985 9 July 1985 
3(2) Nov. 1965 9 Dee. 1965 za) 1986 7 March 1986 
3(3) May 1966 31 May 1966 27(2) 1986 17 Dec. 1986 
4(1) July 1966 18 Aug. 1966 28(1-2) 1988 19 Jan. 1988 
4(2) Aug. 1966 31 Aug. 1966 29(1) 1988 28 June 1988 
5(1) Оес. 1966 31 Dec. 1966 29(2) 1988 16 Dec. 1988 
5(2) June 1967 23 June 1967 30(1-2) 1989 1 Aug. 1989 
5(3) Sept. 1967 30 Sept: 1967 31(1) 1989 29 Dec. 1989 
6(1=2) Dec. 1967 31 Dec. 1967 31(2) 1990 28 May 1990 
6(3) June 1968 30 June 1968 32(1) 11990 30 Nov. 1990 
7(1) Dec. 1968' 31 March 1969 32(2) 1991 7 June 1991 
1(2=3) July 1969 13 Oct. 1969* 33(1-2) 199 6 Sept. 1991 
8(1-2) Oct. 1969 11 Nov. 1969 34(1-2) 1992 9 Sept. 1992 
Sl) Nov. 1969' 16 June 1970 35(1) 1993 14 July 1993 
9(2) Dec. 1969' 20 July 1970 35(2) 1993 2 Dee. 1993 
10(1) May 1970 14 Nov. 1970 36(1-2) 1995 8 Jan. 1995 
10(2) Dec. 1970' 10 July 1971 37(1) 1995 13 Nov. 1995 
11(1) Sept. 1971 8 Oct. 1971 37(2) 1996 8 March 1996 
11(2)° May 1972 21 June 1972* 38(1-2) 1996 17 Dec. 1996 
12(1) 1973 25 July 1973 39 (1-2) 1998 13 May 1998 
12(2) 1973 11 March 1974 40(1-2) 1998 17 Dec. 1998 
1312) 1973 21 Pep. 1973 41(1) 1999 22 Sept. 1999 
14(1—2) 19731 23 Jan. 1974 41(2) 1999 31 Оес. 1999 
15(1) 1975 18 Бес. 1975 42(1-2) 2000 18 Oct. 2000 
15(2) 1978 15 July 1976 43(1-2) 2001 20 Aug. 2001 
16(1) 1977 12 Aug. 1977 44(1) 2002 8 Feb. 2002 
16(2) 1977 17 Sept. 1977 44(2) 2002 30 Aug. 2002 
17(1) 1978 и Feb: 1979 45(1) 2003 29 Aug. 2003 
17(2) 1978 27 July 1978 45(2) 2004 22 March 2004 
18(1-2) 1979 18 May 1979 46(1) 2004 23 Aug. 2004 
19(1) 1979 19 Sept, 1979 46(2) 2004 30 Dec. 2004 
19(2) 1980 14 April 1980 47(1—2) 2005 20 July 2005 
20(1) 1980 22 Aug. 1980 48(1—2) 2006 16 Feb. 2006 
20(2) 1981 17 June 1981 49(1) 2006 10 Nov. 2006 
210 2) 1981 8 Dec. 1981 49(2) 2007 27 July 2007 


Eugene V. Coan 


Department of Invertebrate Zoology and Geology 
California Academy of Sciences, Golden Gate Park, 


1 Five cover dates off by a calendar year. 
2 Issue misdated on back of volume title page as “31 July”; corrected on back of title p. of Vol. 9. 


San Francisco, California 94118-4599, U.S.A.; gene.coan@sierraclub.org 


3 Inside of back cover of Vol. 11(2) listed the dates of previous issues; many subsequent issues included 
partial listings of dates of recent issues as part of index or end-matter. 

4 Day of month of issue not previously indicated; received at The Academy of Natural Sciences, Philadel- 
phia and Library of Congress June 21. 


361 


ee esa, eo ay Be a Ua 
A 5 SRE I FL ee TE : o I <a 
| ur . Sm 10 Lan ne LL eee le u Я Ра 
OS rar ne SE Pen u ee = Er 
A E ze x a а Sa sh" | Ler re iS 2 = ит) у 
RS a | Ser + i. Y Pr А 7 
er 


| em 200% ADOJO9A er 


A ct ever ut 25 ег LU "RS 
gees IE: ZB. | (5500 SR Ever sa 3 el” pd 
IA о мс с а 
GGT лете. о м met EE Erg Der 
0008 yo Bt 45 ‚Bar CE” =." BST 58! ег A ee 
DIE PAE “rote = SPER Ver ist OT. RE ETS ee и 
Dogs 9918 OUT TE ES FAO ПРАХ Maren fee | 
LOUE AA, > 2 eo tome Hi - Tier Be те BETTER or Вос 
сих PORES oP Shes ET oe Maur A ‘Over dat Ti ster ae ANNE 
ODS cee A т BTE vik = CUOMO > забот BEER 0 
RES GA ES LE D: HET veM or ote E BE. 5 
DRG EDER о {Sie - Brut Jone GE.) RS nun (NIGER 
alee. к. IFE OBST nad be ОВ te ra | 
2005 Er LOC © re DORE tea Sc © CBBE es — 
900$ wot Gh BO as В "Вт ani tr TBB: >: Ls ‘S108. by 
TORE win TS 05 4 5 Bet manuka DE | Pht ; shock FS, г. 
Bi oe — 7 o —— mony Fr nan : es > y 


DAA re Se us en 


y р 7 ; BF as => 


La ET mag ss 
Dates sea room or aa pao s 
я ENT а bavi +" fours. "> 


Me o 5 р 7 Fi, | 
u ca je PA Fr Fir: u 2 +h. > Ne a = E 
ej LA, Lo. See 2 et ha ae ME 


MALACOLOGIA, 2008, 50(1-2): 363-392 


INDEX 


Taxa in bold are new; pages in italic indicate 
figures of taxa. 


abbreviata, Cuspidaria 153 
Tropidomya 80, 96, 110, 115-116, 118, 130 
aberrans, Gadila 50 
aberrata, Ledella 130, 151, 153-155, 173 
aberrenta, Ledella 68, 100, 120 
Abra 58, 107 
longicallis 77, 89, 162 
profundorum 77, 86-88, 94, 103-110, 
123, 136, 142-154, 156-157, 165-166, 
168 
abrupta, Rhinoclama 80, 137, 171 
abyssicola, Kelliella 77, 108—110, 153, 160- 
161 
Lyonsiella 80, 89-93, 95, 100, 118, 122- 
123, 157 
abyssorum, Dacrydium 65, 72, 85, 88, 101, 
103-105, 109-110, 125, 127, 142, 144- 
155. 161, 164.169 173 
Lametila 68, 83, 96-97, 99-100, 113, 140, 
142, 144 
Malletia 71, 84, 87-88, 97-105, 110-111, 
119-120, 139, 145-148, 150-155, 158, 
151, 169, 173 
Portlandia 71, 161, 167, 169, 173 
Acanthocardia echinata 77, 122 
acherax, Solemya 90 
acinula, Ledella 68, 83, 106-107, 112, 152, 
154.101. 170 
Acmaeidae 50 
acobambensis, Drymaeus 268 
Acrostoma 332, 334 
Actinonaias 310 
ligamentina 306, 308, 309, 311, 315, 316- 
318 
sapotalensis 312 
acuminata, Hindsia 239 
Ledella 68, 83, 122-123, 137, 141 
Nassaria 239 
Acusta despecta sieboldiana 16, 17, 36, 36, 
56 
acuta, Nuculana 69, 88-89 
Physa 208 
Physella 301 
acutifilos, Juga 202 
adamantina, Tryonia (Paupertryonia) 200 
Adamietta 336 
adamsi, Kelliella 77, 107 
adamsiana, Pisulina 53 
Addisonia lateralis 51 
Adelomelon ferussacii 243 


Adipicola 64 
simpsoni 72, 135 
adusta, Temesa (Neniatracta) cuencaensis 
270 
aeolata, Tindariopsis 68, 107-108 
aequistriata, Gracilinenia 268 
Aeropictus 268 
Aesthenotherus hempilli 
affinis, Limopsis cristata 72, 91-93, 95 
africana, Terebra 243 
Yoldiella inconspicua 70, 84, 159-160, 
162, 160, 172 
africanus, Sinupharus 231 
Solen 231 
Afrophysa 211 
Afropomus balanoudeus 53 
agatheda, Tindaria 83, 156 
Tindariopsis 68, 107-108, 112 
agitata, Pseudoglandina 268, 274 
agueroi, Bostryx (Peronaeus) 268 
Bostryx (Peronaeus) beltrani 268 
aguilari, Bostryx (Bostryx) 268 
alamosae, Tryonia (Paupertryonia) 200 
alaskana, Volutomitra 55 
alba, Pristigloma 67, 87, 95, 97-101, 103, 
111, 113, 119, 137, 141, 146, 152-153, 
157, 167—168, 173 
albicilla, Меты 15, 17, 20, 21, 52 
albicolor, Peruinia 268 
albiconica, Limnaea (Stagnicola) 200 
albilabris, Pupoides (Pupoides) peruvianus 
273 
albocostata, Temesa (Temesa) 268 
Temesa (Temesa) pygmaea 274 
Alcidia 8 
alveus 2 
paivana 6 
Alexandrium 345 
fundyense 344 
alfi, Helminthoglypta 176, 182, 201 
alleni, Thyasira 74, 85, 115, 160, 163-164, 
169-171 
altispira, Euglandina 268 
altorum, Mesembrinus (Mormus) expansus 
268 
Systrophia (Systrophia) 268 
alveus, Alcidia 2 
Artemon 2 
Hypselartemon 1-2, 3, 4, 6, 8-9 
Rectartemon (Hypselartemon) 1-2 
Streptaxis 2 
Streptaxis (Artemon) 2 
Streptaxis (Eustreptaxis) 2 


364 INDEX 


Amalda australis 243 
Amathina triccarinata 56 
Amathinidae 56 
ambiannulatus, Cyclopecten 74, 138, 150, 
152, 164 
Amblema 303-304, 308, 310-312 
plicata 306, 309, 311, 315, 316-318 
Ambleminae 303-305 
Amecanauta 201 
jaliscoensis 201, 205 
americana, Yoldiella 69, 83, 87-88, 100-105, 
110-111, 120, 159 
Amnicolidae 200, 203-204 
Amoria (Amoria) grayi 243 
turneri 243 
Amphibulimidae 268, 274 
Amphimelania 176 
Amphiplica knudseni 51 
Amphora 344 
amphora, Melo 242 
Voluta 242 
Ampularia 299 
gigas 299 
levior 299 
pulchra 233 
Ampullariidae 53, 233, 293, 351 
Amuraplexa 201 
Anachis (Costoanachis) fluctuata 240 
Anatina 255 
ancilla, Pseudoliva 55 
Ancillaria 243 
australis 243 
Ancistrocheirus lesueurii 347 
Andinia (Ehrmanniella) dedicata 270 
(Ehrmanniella) flammulata 268 
Andiniella 268 
andivagus, Naesiotus 268 
angelmaldonadoi, Bostryx (Bostryx) 
modestus 268 
angiportus, Newboldius 268 
angispira, Bostryx (Bostryx) obliquiportus 
268 
angolensis, Cetoconcha 78 
angulare, Dacrydium 72, 85, 167 
angularis, Myonera 80, 101, 138, 140, 142, 
154 
angulobasis, Drymaeus 273 
angustus, Bulimulus (Bulimulus) vesicalis 
268 
Annatina 255 
Anodon georginae 229 
susannae 229 
tenuis 229 
Anodontites (Anodontites) trigonus 
georginae 229 
(Anodontites) exoticus susannae 229 


Anomiidae 59, 72, 121 
antarctica, Laevipilina 49 
Anula 256 
Apachecoccus 201 

arizonae 201 
apinensis, Glycemeris 230 

Glycymeris 228, 230 
Aplexa 201 

microstriata 211 
Aplexinae 201, 211 
Aplysia 45 

oculifera 16, 32, 33, 56 


_Aplysiidae 16, 32, 33, 41, 45, 56 


Appisania 240 
aquatilis, Haliotis diversicolor 15, 17, 19, 19, 


Sulculus diversicolor 51 
araozi, Bulimulus (Bulimulus) 268 
arboriferus, Neopetraeus obesus 273 
Neopetraeus paucistrigatus 273 
Archiphysa 201 
ashmuni 201 
sonomae 210 
Architectonicidae 55 
Arcidae 59, 65 
argellacea, Helix 245 
argentata, Philine 16, 31, 32, 56 
argentinae, Ledella pustulosa 69, 118 
argentinea, Ledella pustulosa 83, 117-120 
Yoldiella 115-115 
argentineae, Limea 73, 115 
argentinensis, Yoldiella 69 
argentinus, Illex 347, 348, 349 
Pupoides (Ischnopupides) chordatus 268 
argilacea, Helix 245 
argillacea, Helix 245 
Arionidae 182, 203, 208 
arizonae, Apachecoccus 201 
arntzi, Micropilina 49 
Artemon 1-2, 8 
alveus 2 
contusulus 4 
deshayesianus 5 
paivanus 6 
articulata, Cyclostoma 237 
Tropidophora 237 
artipica, Yoldiella 69, 162, 164, 166 
asellus, Lepidopleurus 49 
ashmuni, Archiphysa 201 
Aspergillium 256 
Aspergillum 256 
asperula, Bentharca 71, 84, 95, 98-99, 101- 
102, 104-105, 109, 113, 152, 163, 167- 
169, 172-173 
Assiminea 41 
cienegensis 209 


INDEX 


Japonica 15, 24, 26, 54 
pecos 209 
Assimineidae 15, 24, 26, 54, 209 
Astarte 77, 86, 88-90, 102, 121, 176 
triangularis 88 
Astartidae 59 
Asterophila japonica 54 
Asthenotherus hemphilli 78, 114 
atacellana, Deminucula 67, 82, 87, 90-100, 
106-108, 117-119, 123-132, 134-137, 
139-141, 143, 149, 156, 162-165, 169 
atlanta, Halonympha 80, 88, 97-98, 123, 
137, 166 
atlantica, Bidentaria 100, 156 
Cuspidaria 79, 90, 95, 105-106, 140, 153, 
156, 171 
Halonympha 96 
Kelliella 77, 86, 90, 97, 106-112, 117- 
120, 122, 124-133, 135-157, 159-160, 
163, 165-166, 168-170, 172-173 
Malletia succisa 105 
Myonera 80, 86-88, 92, 101, 109, 111, 
16117, 126; 140,147, 1494155458, 
173 
Policordia 81, 95, 128-129, 131-132, 138, 
147, 158, 160, 164 
Protocuspidaria 78, 92 
Thyasira 74, 104, 145, 152 
Thyasira subovata 116 
Thyasira succisa 75, 86, 89, 92, 96, 106- 
108, 115, 41729123. 152416365: 
170,172 
aurantiaca, Melanodrymia 52 
Auricula (Chilina) fluctuosa 321 
auriculare, Cyclostoma 233 
aurisvulpina, Bulimus 244 
Chilonopsis 244 
auris-vulpina, Bulimus 244 
Voluta 244 
aurita, Limopsis 72, 85, 90, 121, 124 
australis, Amalda 243 
Ancillaria 243 
Austrinauta 201 
Austrodiscus superbus tucumanus 272 
Austroselenites weyrauchi 276 
Aximea 256 
Axinaea 256 
Axinodon symmetros 76, 112, 125-127, 131, 
138, 142, 146 
Axinulus incrassatus 129-134, 137-139, 
143 
Axinus grandis 74, 98, 161 
Aylacostoma (Hemisinus) lineolata 235 


bacıllus, Hastula 55 
Bakerilymnaea 268 


365 


Balanocochlis 331-332, 334, 336 
glandiformis 334 
glans 331-335, 333-334, 336, 337 
pisum 333 
balanoudeus, Afropomus 53 
bambamarcaénsis, Naesiotus (Naesiotus) 
268 
barnardi, Cuspidaria 79, 95, 117, 161 
basiplanata, Epiphragmophora 268 
Bathyacmaea 43 
secunda 50 
Bathnyarca IVAR AS 
glacialis 71, 89 
inaequisculpta 71, 84, 99, 101, 108-111, 
113, 136, 140-142, 144, 149-152, 154, 
158-161, 165-169 
pectunculoides 71, 85, 88-92, 109, 121, 
156, 169 
pectunculoides pellucida 170 
Bathypecten 73, 85, 93, 107, 112-113, 115- 
118, 123-124, 150, 158, 172 
eucymatus 73, 87, 91-95, 109, 118-119, 
123, 140 
Bathyphytophilidae 51 
Bathyphytophilus 43 
diegensis 51 
Batillaria californica 202 
eumingi1d, 22-23, 58 
Batillariidae 15, 22, 23, 53 
Belgrandiinae 204 
beltrani, Bostryx (Peronaeus) agueroi 268 
Bentharca asperula 71, 84, 95, 98-99, 101- 
102, 104-105, 109, 113, 152, 163, 167- 
169, 172-173 
nodulosa 71, 157 
bequaerti, Drymaeus 268, 272 
Temesa (Neniatracta) 268 
Bequaertinenia 268 
bermudezae, Bostryx (Pseudoperonaeus) 
268, 274 
bernardinus, Yaquicoccus 201, 211 
beyrichii, Midotrochus 52 
bicarinatus, Trochus 232 
bicolor, Bulinus 275 
Naesiotus (Naesiotus) 269 
Bidentaria atlantica 100, 156 
bifasciata, Natica 236 
Polinices (Polinices) 236 
biguttata, Yoldiella 69, 106-108, 112, 117 
Bilamelliferus 269 
bilanta, Yoldiella 69, 83, 158-160, 162, 167, 
170—172 
billenheusti, Phos 238 
Biomphalaria kansasensis 209 
birabenorum, Bostryx (Lissoacme) 269 
biscayensis, Kelliella 77, 86, 121-123 


366 INDEX 


Thyasira 74, 109, 152, 155-156, 172 
Yoldiella 70, 83, 132, 134, 136, 139-140, 
142-155 
bisecta, Limatula 72, 121 
bitubercularis, Nassaria 239 
blakeana, Pyrgulopsis 201 
blanda, Yoldiella 69, 83, 117-120 
boivinii, Cypraea 15, 17, 23, 24, 44, 53 
borealis, Olivella 55 
Bostryx 270-271, 273-274 
(Bostryx) aguilari 268 
(Bostryx) bromeliarum grandiportus 271 
(Bostryx) chusgonensis 269 
(Bostryx) haasi 271 | 
(Bostryx) haasi minor 272 
(Bostryx) modestus angelmaldonadoi 268 
(Bostryx) obliquiportus 273 
(Bostryx) obliquiportus angispira 268 
(Bostryx) obliquiportus inflatiportus 271 
(Bostryx) obliquiportus laraosensis 272 
(Bostryx) ortizi 273 
(Bostryx) pygmaeus 274 
(Bostryx) pygmaeus costatus 269 
(Bostryx) rodriguezae 274 
(Bostryx) scotophilus 274 
(Bostryx) vilchezi 275 
(Bostryx) willinki 275 
(Bostryx) zilchi 275 
(Bostryx) zilchi compactus 269 
(Bostryx) zilchi glomeratus 271 
cunyacensis 270 
(Elatibostryx) imeldae 270-271 
(Elatibostryx) imeldae costifer 269 
(Elatibostryx) rehderi 274 
florezi 270 
(Lissoacme) birabenorum 269 
(Lissoacme) globosus 271 
(Multifasciatus) Superbus 275 
(Peronaeus) адиего! 268 
(Peronaeus) адиего! beltrani 268 
(Peronaeus) crucilineatus 270 
(Platybostryx) weyrauchi 273 


(Pseudoperonaeus) bermudezae 268, 274 


(Pseudoperonaeus) cylindricus 270 
(Pseudoperonaeus) lizarasoae 272 
(Pseudoperonaeus) longispira 272 
(Scansiocohlea) gracilis 271 
weyrauchi 276 
Bradybaenidae 16, 17, 36, 36, 56 
braziliensis, Cetoconcha 78, 114 
brevicula, Littorina 15, 17, 23, 25, 53 
Brevinucula subtrangularis 67, 112 
verrilli 67, 82, 95, 97, 99-100, 102, 107- 
109, 113, 137, 140, 142, 152, 156-163 
brevis, Thyasira 74, 85, 87, 93-94, 97-101, 
104, 117-119, 124-127, 129-135, 137- 


139, 141-142, 145, 147, 149-153, 155, 
162, 164—168, 172 
bridgesii, Ротасеа 353, 354 
broderipi, Voluta 242 
broderipii, Melo 222 
Melo (Melocorona) 242 
Voluta 242 
broderippii, Voluta 242 
bromeliarum, Bostryx (Bostryx) grandiportus 
| 
Brontes 256 
Brontis 256 
Brotia 234, 332, 336 
carolinae 233 
costula 233, 332 
hainanensis 300 
henriettae 234 
testudinaria 337 
brunei, Tryonia (Paupertryonia) 202 
Buccinidae 16, 27, 28, 54, 228, 238 
Buccinum fasciculatum 240 
laevissimum 256 
pristis 239 
undosum 238 
buckleyi, Elliptio 304 
Popenaias 304 
bulbosa, Melania 208 
Bulimulidae 8, 208-209 
Bulimulus (Bulimulus) araozi 268 
(Bulimulus) vesicalis angustus 268 
(Protoglyptus) sarcochrous 275 
(Rhinus) thomei 275 
Bulimus aurisvulpina 244 
auris-vulpina 244 
subroseus 273 
tschudii 269 
veranyi 268 
Bulinus bicolor 275 
Bullaea 227 
semiplicata 241 
Bullia 227, 241 
semiplicata 227, 241 
burchardi, Mitrella 54 
bushae, Nuculoidea 67, 82, 88-90, 106, 
122-130, 132-133, 135, 156, 158-159, 
161, 163, 165-169 
Thyasira 74, 163 
Bushia 78, 135 


cahuillarum, Pyrgulopsis 202 

cajamarcana, Steeriana (Steeriana) 269 

Calibasis 202 

californica, Batillaria 202 

californiensis, Fontelicella 185, 203 
Fontelicella (Fontelicella) 202 

callicredemma, Nucula 67 


INDEX 367 


Nuculoma 113 
Calliostoma (Maurea) selectum 232 
Calliostomatidae 232 
Calliotropis 233 
callistiformis, Tindaria 68, 83, 98-105, 109- 
141.147.119. 126, 130, 14831441538) 
161, 164, 168-169 
Calliteuthis dofleini 349 
Calotropis 233 
Calyptraeidae 15, 17, 23, 23, 45, 53 
camachoi, Neopetraeus 269 
Camaenidae 244, 265 
Campanile 236 
symbolicum 236 
Campanilidae 236 
campbelli, Strombus 237-238 
Strombus (Doxander) vittatus 238 
canaliculata, Pomacea 293-294, 295, 296- 
297, 298, 299-301, 351, 352-353, 354— 
355 
Cantharus (Gemophos) elegans 238 
(Gemophos) vibex 239 
(Prodotia) iostoma 238 
capensis, Yoldiella 70, 83, 164, 167, 170- 
17% 
caramba, Paludiscala 188, 202, 208 
Cardiidae 59, 78, 114 
Cardiomya 100, 111, 118 
costellata 79, 96; 106-121,123,:137; 141, 
153 
curta 79, 92 
gemma 116 
knudseni 79, 89, 96-98, 100, 115-117, 128 
Carditid 86 
Carditidae 59, 77, 88, 114-115, 121, 123 
Caribnauta 201-202 
harryi 202, 204 
carinata, Pleurotoma 243-244 
Pleurotoma decussata 244 
Carinifex shotwelli 210 
carinifex, Menetus 202 
carmelita, Helix 245 
carolinae, Brotia 233 
Melania 233 
carpenteri, Propeleda 69 
Propoleda 83, 114-116 
carranzae, Mexipyrgus 202-203, 206-207 


carrozae, Thyasira 74, 85, 94, 115-117, 150, 


158, 163-165, 170-171 
Caryodidae 244 
Cavolinia uncinata 16, 32, 33, 56 
Cavoliniidae 16, 32, 33, 41, 45, 56 
ccharopa, Microdiscula 55 
cedis, Incertae 81-82, 87 

Incerte 87-90, 94-99, 102-104, 107-108, 

110-111, 114-117, 119, 122-125, 128, 


130, 136-138, 141, 143, 145, 147, 150, 
153, 160-161, 164-165, 169-173 
celendinensis, Drymaeus 269 
Steeriana (Steeriana) 269 
Steeriana (Steeriana) isidroensis 272 
Steeriana (Steeriana) minor 272 
Cellana 42 
grata 15-16, 17-18, 50 
toreuma 50 
celtica, Limatula 72, 105, 114, 142, 147, 150, 
152 
centobi, Propeamussium 74, 157 
Ceratoptilus 236 
Cerithidea 197, 235 
obtusa 235 
reidi 235-236 
Cerithioidea 331 
Cerithium laeve 236 
laevis 236 
leve 236 
reidi 236 
rodeoensis 202 
truncatum 235-236 
cerrateae, Epiphragmophora (Karlschmidtia) 
269 
Hemicena 269 
Naesiotus (Raphiellus) 269 
Cetoconcha 78, 143 
angolensis 78 
braziliensis 78, 114 
chamayensis, Naesiotus (Naesiotus) 
subcostatus 269 
championi, Pseudotindaria 68, 117, 119- 
120,152 
Charopidae 265, 270-272, 274-276 
chauliodonta, Gastrocopta (Gastrocopta) 
202 
cheatumi, Potamopyrgus 209 
Chiapaphysa 202, 209 
grijalvae 202, 204 
расйса 208 
Chicoracea 254 
Chicoreus 254 
childreni, Unio 229-230 
chilensis, Diplodon (Diplodon) 230 
Unio 230 
chiletensis, Scutalus (Scutalus) 269 
Scutalus (Scutalus) granulatus 271 
Chilina 321, 328 
gallardoi 328 
guaraniana 328 
iguazuensis 321-322, 322-324, 324— 
326, 326-328, 328-329 
megastoma 321, 323, 327-329, 329 
parva 329 
Chilinidae 321 


368 | INDEX 


Chilonopsis aurisvulpina 244 
chinensis, Cipangopaludina 233 
Cipangopaludina laeta 15, 17, 22, 22, 53 
Cuspidaria 232 
Glauconome 230 
Mitra 242 
Neaera 228, 232 
Neroea 232 
Paludina 233 
Unio 230 
Viviparus 233 
chippevarum, Laurentiphysa 202 
Chiton 256 | 
squamosus 256 
Chlamys (Azumapecten) farreri farreri 279, 
289 
farreri farreri 279-281, 282-284, 286-288, 
289-291 
Chlorostroma lischkei 52 
Chonarus 255 
Chondrus 255 
chordatus, Pupoides (Ischnopupides) 
argentinus 268 
Choristella 43 
hickmanae 51 
Choristellidae 51 
Chromodorididae 16, 32, 34, 56 
Chrondrus 255 
chrysopylica, Juga 202, 205 
chupaderae, Fontelicella 203 
churinceanus, Mexipyrgus 202-203 
chusgonensis, Bostryx (Bostryx) 269 
cienegensis, Assiminea 209 
Ciliatocardium ciliatum 122 
ciliatum, Ciliatocardium 122 
Clinocardium 78 
Cinnalepeta pulchella 53 
Cipangopaludina chinensis 233 
chinensis laeta 15, 17, 22, 22, 53 
circinata, Cuspidaria 79, 113, 117, 120 
circumstriata, Iryonia 211 
cisternina, Physa 206 
Cittarium 255 
clarkei, Prelametila 68, 83, 97, 108, 119-120 
Clausiliidae 265, 268-270, 272-276 
Clavatula griffithii 222, 244 
clavatum, Pseudamussium 74, 121 
clavella, Pyrula 241 
clavigera, Thais 16, 17, 29, 30, 54 
cleliae, Zilchogyra 269 
Clenchiella 203 
Clinocardium ciliatum 78 
Clypeosectus delectus 51 
coahuilae, Durangonella 203 
Coahuilix 203-204 
hubbsi 203, 205 


Coccopygya hispida 52 
Cocculina 39 
japonica 15, 20, 21, 52 
nipponica 52 
Cocculinella 43 
minutissima 51 
Cocculinellidae 45, 51 
Cocculinidae 15, 20, 27, 39, 52 
cochisi, Pyrgulopsis 201 
Cochliopidae 200, 271, 273-274 
Cochliopina milleri 207 
Cochliopinae 203, 206-209 
Cochlodesma tenerum 78, 122, 128, 131, 160 
coelestini, Drymaeus (Diaphanomormus) 
obesus 270, 273 
Coleoidea 349 
Columbella fluctuata 240 
suturalis 240 
tylerae 240 
Columbellidae 16, 28, 28, 54, 240 
combinai, Mesembrinus (Ornatimormus) 269 
Columbinia 274 
(Pfeifferiella) haasi 271, 274 
(Pfeifferiella) subterranea 275 
columellaris, Naesiotus (Naesiotellus) 273 
commutata, Nuculana 69, 120 
compactus, Bostryx (Bostryx) zilchi 269 
comptus, Ischnochiton 15, 40, 49 
concentrica, Kelliella 77, 90-91 
Rhynchopelta 52 
conchos, Disconaias 203 
confluentis, Pyrgulopsis 211 
confusa, Monodonta labio 19, 20, 52 
Monodonta labio forma 15 
conica, Hipponix 15, 25, 27, 54 
Melania 234 
Paludomus 234 
Pila 351, 352-353, 354-355 
Conidae 16, 30, 55 
conradi, Thracia 78, 89, 91 
constrictus, Sinonovacula 230 
Solen 230 
contermina, Helix 3, 4 
contusula, Helix 1-2, 5 
Helix (Helicogena) 4 
contusulus, Artemon 4 
Hypselartemon 1-2, 3, 4-5, 9 
Conus ebraeus 16, 30, 55 
coraeformis, Scutalus (Scutalus) 
debilisculptus 270 
Corbicula 229, 355 
Corbiculidae 196, 228 
Corbula 78, 92, 121 
Cornirostra pellucida 56 
Cornirostridae 56 
cornuarietis, Marisa 53 


INDEX 369 


corpulenta, Neilonella 68, 113 
corrugata, Cuspidaria 232 
costata, Siliqua 231 
Costatella 209 
costatus, Bostryx (Bostryx) pygmaeus 269 
Leguminaria 231 
Solen 231 
costellata, Cardiomya 79, 96, 106, 121, 123, 
137, 141, 1683 
costifer, Bostryx (Elatibostryx) imeldae 269 
costula, Brotia 233, 332 
Melania 233 
costulatus, Scutalus (Vermiculatus) 269 
Crassatella cuneata 231 
ornata 230 
Crassatellidae 230 
crassicostata, Temesa (Temesa) decimvolvis 
269 
Crassostrea gigas 290 
virginica 290, 344 
Craterarion 203 
pachyostracon 182, 203, 208 
crenilabrum, Pisania 240 
crenulatus, Lopesianus 270, 272 
Crepidula onyx 15, 17, 23, 23, 53 
crispa, Pleurostoma 244 
Turris 244 
Cristaria (Pletholophus) discoidea discoidea 
229 
cristata, Limopsis 85, 89-90, 94, 111, 116, 
123, 138, 164, 170 
Limopsis affinis72, 91-93, 95 
Limopsis cristata 72, 123, 125, 156 
Limopsis lanceolata 72, 164, 171 
Limopsis intermedia 72, 106-107 
crossei, Streptaxis 6 
croulinensis, Thyasira 74, 85, 88-98, 105- 
106, 114-116, 119, 121-122, 124, 136- 
138, 142, 144, 147, 152, 159-161, 163- 
166, 170-172 
crucilineatus, Bostryx (Peronaeus) 270 
Cryptosoma javanica 237 
cuencaensis, Temesa (Neniatracta) adusta 
270 
culmineus, Scutalus (Vermiculatus) zilchi 275 
cumingii, Batillaria 15, 22, 23, 53 
cuneata, Crassatella 231 
Malletia 71, 84, 87, 96-102, 116, 118-119, 
127-128, 134-136, 138, 140-151, 153- 
167, 182 
cunninghami, Helix 244 
Trochus 232-233 
cunyacensis, Bostryx 270 
curta, Cardiomya 79, 92 
Cuspidaria 96 
Yoldiella 70, 83, 91-95, 106-107, 112, 


115-116, 123-132, 134-136, 158-159, 
164, 166-167, 171-172 
Cuspidaria 79, 86-87, 89-91, 94-96, 99- 
101, 103, 105-107, 110-111, 113, 115- 
116, 118-122, 131, 133-135, 138-140, 
143-144, 146-147, 149, 151, 153-157, 
161, 165-167, 228, 232 
abbreviata 153 
atlantica 79, 90, 95, 105-106, 140, 153, 
156, 171 
barnardi 79, 95, 117, 161 
chinensis 232 
circinata 79, 113, 117, 120 
corrugata 232 
curta 96 
cuspidata 121 
inflata 88, 98 
jeffreysi 79, 93, 133, 156 
obesa 79, 86, 91, 95, 108, 128, 130-131, 
133-134, 139-140, 143, 147 
parva 79, 86-96, 99, 106, 113, 118, 122- 
123, 125-142, 144-150, 153, 156-158, 
160, 166 
rostrata 92 
teres 157 
undata 79, 103 
ventricosa 79, 163 
Cuspidariidae 58-59, 62, 228, 232 
cuspidata, Cuspidaria 121 
cuzcoensis, Scutalus (Vermiculatus) 270 
Scutalus (Vermiculatus) marasensis 272 
Cyclopecten 74, 85, 92, 96, 105, 110, 113, 
116, 119, 123-124, 126, 142, 150, 158- 
161,163, TOP/109, 171 
ambiannulatus 74, 138, 150, 152, 164 
pustulosus 74, 89, 106, 110, 118 
simplex 74, 92, 105 
Cyclophoridae 15, 17; 21, 22 53. 233 
Cyclophorus herklotsi 15, 21, 22, 53 
Cyclostoma 237 
articulata 237 
auriculare 233 
madagascariensis 237 
pulchra 237 
pulchrum 237 
Cylichnidae 16, 31, 33, 41, 45, 56 
cylindricus, Bostryx (Pseudoperonaeus) 270 
Cyllene 228, 241 
owenii 222, 228, 240-241 
Cypraea boivinii 15, 17, 23, 24, 44, 53 
gracilis japonica 44 
Cypraeidae 15, 17, 23, 24, 39, 44, 53 
cyprinoides, Cyrena 228 
Cyprogenia 304 
Cyrena cyprinoides 228 
similis 228-229 


370 INDEX 


Cyrtonaias 304, 310 


tampicoensis 306, 309, 311, 315, 316-318 


Cytherea dione 255 
dronea 255 
dronia 255 


Dacryaium 58, 65, 12111, 1155122, 138. 
173 
abyssorum 65, 72, 85, 88, 101, 103-105, 
109-110, 125, 127, 142, 144-155, 161, 
164, 169, 173 
angulare 72, 85, 167 
hedleyi 72, 109-110 
ockelmanni 72, 85, 91-95, 115-117, 123- 
135, 137, 139-140, 156, 158, 160-161 
sandersi 72, 85, 92, 96-98, 112-114, 
138-139, 143 
vitreum 72, 88-89 
wareni 72, 92, 123, 135 
Darina 232 
solenoides 232 
Daudebardiaella 204 
davisi, Fontelicella 203 
deaurata, Mactra 231 
deauratum, Mesodesma 231 
debilisculptus, Scutalus (Scutalus) 
coraeformis 270 
Thaumastus (Thaumastiella) occidentalis 
270 
decimvolvis, Temesa (Temesa) 270 
Temesa (Temesa) crassicostata 269 
Temesa (Temesa) mantaroensis 272 
Temesa (Temesa) minor 272 
decussata, Pleurotoma carinata 244 
dedicata, Andinia (Ehrmanniella) 270 
dedleyi, Peculator 55 
deformis, Strombus 238 
Delectopecten 73, 85, 122-123, 136 
vitreus 73, 85, 121, 123, 136 
delectus, Clypeosectus 51 
Demarestia firoloida 254 
demarestia, Firoloida 254 
Deminucula atacellana 67, 82, 87, 90-100, 
106-108, 117-119, 123-132, 134-137, 
139-141, 143, 149, 156, 162-165, 169 
demistriata, Myonera 80, 92, 96, 98-99, 119, 
141,153 
densestrigatus, Mesembrinus (Ornatimormus) 
henrypilsbryi 270 
densicostata, Policordia 81, 91-93, 99, 136- 
137, 171-172 
Dentaliidae 15, 40, 50 
Dentalium octanglatum 15, 40, 50 
denticulata, Erycina 231 
Mactra 231 
Mesodesma 231 


depressa, Halonympha 80, 96, 123, 134, 
153,157 
deshayesianus, Artemon 5 
Hypselartemon 1-2, 3, 4-6, 8-9 
Streptaxis 5-6 
Streptaxis (Eustreptaxis) 5 
despecta, Acusta sieboldiana 16, 17, 36, 36, 
56 
diagonalis, Tropidomya 80, 164 
Diaphanomormus 270 
didyma, Glossaulax 15, 24, 25, 54 
diegensis, Bathyphytophilus 51 
diffusa, Pomacea 299, 351, 353, 354 
diluta, Epiphragmophora semiaperta 274 
Epiphragmophora semiclausa 274 
dineana, Limnaea (Pseudosuccinea) 203 
dione, Cytherea 255 
Diplodon (Diplodon) chilensis 230 
Dipsada 256 
Dipsas 256 
discoidea, Cristaria (Pletholophus) discoidea 
229 
Symphynota 229 
Disconaias conchos 203 
discors, Musculus 72, 127 
dissimilis, Yoldiella 70, 87, 99, 138, 141 
diversicolor, Haliotis aquatilis 15, 17, 19, 19, 
5 
Sulculus aquatilis 51 
dofleini, Calliteuthis 349 
dolioides, Pomacea 353 
dombeyanus, Plectomerus 306, 309, 311, 
315, 316-318 
domesticus, Passer 355 
douglasiae, Unio 229 
Dreissena polymorpha 351 
Dromus 304 
dronea, Cytherea 255 
dronia, Cytherea 255 
Drymaeus 270 
acobambensis 268 
angulobasis 273 
bequaerti 268, 272 
celendinensis 269 
(Diaphanomormus) coelestini obesus 270, 
278 
(Drymaeus) souzalopesi 274 
(Drymaeus) translucidus 275 
(Mesembrinus) pseudobesus 270 
(Mormus) expansus flavilabrum 270 
(Ornatimorus) multiguttatus 273 
(Orodrymaeus) farrisi quadritaeniatus 274 
pilsbryi 274 
pseudobesus 273 
sulfureus obesus 273 


Durangonella 203, 205 


INDEX 371 


coahuilae 203 
durouchouxi, Thracia 78, 106 


ebena, Fusconaia 312 
ebraeus, Conus 16, 30, 55 
echinata, Acanthocardia 77, 122 
Edentaria simplis 103, 153 
Edentulae 5 
edulis, Mytilus 290-291 
Eglisia tricarinata 238 
elata, Physa 201 
Elatibostryx 270 
elegans, Cantharus (Gemophos) 238 
Pusio 238 | 
Triton 228, 238 
Triton (Pusio) 228, 238 
elegantulus, Naesiotus 270 
eliseoduartei, Systrophia (Scolodonta) 270 
ella, Yoldiella 70, 84, 87, 96-100, 108, 113, 
118, 129, 132, 142-150, 152, 154-155, 
161, 163, 167-168 
ellipsiformis, Venustaconcha 306, 309, 311, 
315, 316-318 
elliptica, Tridonta 77, 121-122 
Elliptio 304 
buckleyi 304 
elongata, Kelliella 77, 90, 106, 111, 115-116, 
118—119, 146, 159, 170-171 
Nuculoma 67, 105 
elpis, Epilepton 76, 103 
emarginatius, Gorgoleptis 52 
Emmericiinae 204 
enata, Yoldiella 70, 84, 87, 91, 108 
Endodonta superba 275 
Endodontidae 265, 275 
Enteroctopus megalocyathus 347 
Epilepton 76, 108, 141, 163, 165-166 
elpis 76, 103 
subtrigonum 76, 138 
Epiphragmophora basiplanata 268 
diluta semiaperta 274 
diluta semiclausa 274 
granulosa 271 
(Karlschmidtia) cerrateae 269 
mirabilis 273 
огтеа! 273 
zilchi 275 
Epiphragmophoridae 268-269, 271, 273-275 
Episiphon subrectum 15, 40, 50 
Epitoniidae 238 
equalis, Thyasira 74, 87, 89-94, 97, 106, 
114-115, 117-119, 122-127, 130-131, 
135, 139, 141, 144-146, 148, 154-155, 
158-159, 161-164, 166, 170, 172 
erebus, Pseudotindaria 68, 83, 100, 109- 
110, 113, 118, 154, 161-162, 166-167 


Erycina denticulata 231 
solenoides 231-232 
Subangulata 231 
escobedae, Mexipyrgus 203 
Eucrassatella ornata 230 
eucymatus, Bathypecten 73, 87, 91-95, 109, 
118-119, 123, 140 
Euglandina altispira 268 
rosea 351 
Eulimidae 44, 54 
eumyaria, Thyasira 75, 85, 116, 122-124, 
158, 160,182, 165171 
Euplica scripta 16, 28, 28, 54 
euripes, Temnocinclis 52 
Eustreptaxis 5 
ewingi, Vema 49 
excavata, Thyasira plicata 75, 164 
exoticus, Anodontites (Anodontites) 
susannae 229 
expansus, Drymaeus (Mormus) flavilabrum 
270 
Mesembrinus (Mormus) altorum 268 
Mesembrinus (Mormus) orcesi 273 
extensa, Yoldiella 70, 84, 117 


fabula, Yoldiella 70, 96-103, 107-109, 119- 
120, 129, 136, 141-143, 145, 148-149, 
151, 155, 162-163 

farreri, Chlamys (Azumapecten) farreri 279, 

289 
Chlamys farreri 279-281, 282-284, 286- 
288, 289-291 

farrisi, Drymaeus (Orodrymaeus) 
quadritaeniatus 274 

fasciculatum, Buccinum 240 
Pisania 240 

Fasciolariidae 240 

fasciolaris, Ptychobranchus 306, 309, 311, 
315, 316-318 

fernandezae, Naesiotus (Maranhoniellus) 
270 

ferrea, Japeuthria 16, 27, 28, 54 

Ferrissia fragilis 355 

ferruginea, Thyasira 75, 85, 87-99, 101, 
103, 105-108, 114-122, 124, 126-127, 
131-132, 135-138, 140, 143-148, 150- 
152, 159-161, 165-166, 172 

ferussacii, Adelomelon 243 
Voluta 243 

Ferussaciidae 244 

festiva, Hypselodoris 16, 32, 35, 56 
Nassarius 16, 29, 29, 54 

fibula, Yoldiella 146, 152, 167-169 

filatovae, Silicula 71, 84, 87, 96, 100, 102- 

105, 111, 123, 143-144, 146, 148-152, 
161, 166, 168-170 


372 


Spinula 69, 83, 124, 126, 158, 160, 162, 
164 
filosa, Lucinoma 74, 88, 164, 169, 171-172 
Firoloida demarestia 254 
firoloida, Demarestia 254 
Fissurellidae 51 
flachi, Peruinia superba 275 
flammulata, Andinia (Ehrmanniella) 268 
flava, Fusconaia 306, 309, 311, 315, 316- 
318 
flavilabrum, Drymaeus (Mormus) expansus 
270 
Flexipecten proteus 73 
Flexopecten proteus 121 
flexuosa, Halicardia 81, 171 
Scutellastra 50 
florezi, Bostryx 270 
Floreziellus 270 
Happia (Happia) 270 
Incania 270 
Phenacotaxus (Ataxellus) 270 
Radiodiscus 270 
Systrophia (Scolodonta) 270 
Floreziellus 271 
florezi 270-271 
fluctuata, Anachis (Costoanachis) fluctuata 
240 
Columbella 240 
fluctuosa, Auricula (Chilina) 321 
Fluminicola junturae 205 
Fontelicella 185, 203-204, 207 
californiensis 185, 203 
chupaderae 203 
davisi 203 
(Fontelicella) californiensis 202 
gilae 204 
kolobensis 205 
metcalfi 206 
(Microamnicola) micrococcus 207 
(Natricola) melina 206 
(Natricola) robusta 207 
pecosensis 209 
pinetorum 205, 209 
roswellensis 210 
thermalis 211 
trivialis 211 
Fontigens 204 
Fontigentinae 204 
forma, Monodonta labio confusa 15 
formosa, Lyonsiella 80, 94, 113-114, 122, 
156-158 
fragilis, Ferrissia 355 
Lyonsiella 80, 91-94, 100 
Silicula 71, 97, 99-100, 118, 126, 128, 133, 
138, 141, 165 
franzenae, Gastrocopta 204 


INDEX 


franzi, Zilchogyra 271 
fraseri, Helix 244-245 

Sphaerospira 245 
freethii, Melania 234 

Potadoma 234 
freilei, Lyonsiella 80, 158, 167 
frethii, Melania 234 
fretterae, Neomphalus 52 
Friersonia 304 

iridella 312 
frigida, Yoldiella 70, 84, 89-91, 108, 121 
fontinalis, Physa 207 
fundyense, Alexandrium 344 
Fusconaia 310 

ebena 312 

flava 306, 309, 311, 315, 316-318 
Fusitriton 42 

oregonensis 15, 25, 26, 54 
Fusus Striata 241 

Striatus 241 


Gadila aberrans 50 
Gadilidae 50 
Gadilinidae 15, 40, 50 
galathea, Ledella 154 
Limopsis 72, 85, 87, 99-103, 105, 109- 
110, 114, 152, 154, 161, 163, 166, 168- 
1691173 
galatheae, Ledella 68, 154, 163 
Limopsis 169 
Neopilina 49 
Galeommatoidea 170 
gallardoi, Chilina 321, 328 
gangetica, Novaculina 231 
Gastrocopta franzenae 204 
(Gastrocopta) chauliodonta 202 
(Gastrocopta) lavernensis 205 
(Gastrocopta) scaevoscala 210 
Geloina 229 
Gemma 176 
gemma, Cardiomya 116 
Policordia 81, 99-101, 113, 118-119, 
125-126, 128, 130, 132-135, 139, 143, 
145-147, 172 
Gemmula speciosa 244 
gentilis, Lutrilimnea 204 
geophilus, Naesiotus (Naesiotus) 271 
georginae, Anodon 229 
Anodontites (Anodontites) trigonus 229 
Melo 222 
Voluta 242 
Gibbula 232 
magus 232 
giganteus, Radiodiscus 271 
gigas, Ampullaria 299 
Crassostrea 290 


INDEX So 


Pomacea 299 
gilae, Fontelicella 204 
Tryonia 204 
glacialis, Bathyarca 71, 89 
Glacidorbidae 55 
Glacidorbis hedleyi 55 
glandiformis, Balanocochlis 334 
glans, Balanocochlis 331-335, 333-334, 
390,397 
Melania 333 
Glauconome 228 
chinensis 230 
Glauconomidae 230 
Glebula 310 
rotundata 306, 309, 311, 315, 316-318 
globosus, Bostryx (Lissoacme) 271 
globulosa, Melania 234 
Paludomus 234 
glomeratus, Bostryx (Bostryx) zilchi 271 
Glossaulax 42 
didyma 15, 24, 25, 54 
Glycemeris 227-228 
apinensis 230 
Glycimeris 227 
Glycymerididae 228 
Glycymeris 227-228 
apinensis 228, 230 
Gocea 204 
golbachi, Radiodiscus 271 
Goniobasis rodeoensis 202 
Goodallia triangularis 77 
Gorgoleptis emarginatius 52 
Gracilinenia aequistriata 268 
gracilis, Bostryx (Scansiocohlea) 271 
Cypraea japonica 44 
Thracia 78, 122 
gracillimus, Naesiotus 271 
grandiportus, Bostryx (Bostryx) bromeliarum 
271 
Zilchiella 271 
grandis, Axinus 74, 98, 161 
Pleurostoma 244 
Pleurotoma 244 
Solemya 67, 91-92, 99 
grandiventris, Scutalus 271 
granulata, Poromya 78, 110, 138 
granulatus, Scutalus (Scutalus) chiletensis 
274 
granulosa, Epiphragmophora 271 
Nuculoma 67, 82, 90-95, 108, 123, 136, 
141 
granulose, Nuculoma 164, 170 
grasslei, Malletia 71, 105-106, 120 
grata, Cellana 15-16, 17-18, 50 
grayanus, Trochus 232 
grayi, Amoria (Amoria) 243 


griffithii, Clavatula 222, 244 
Ptychobela 244 

grijalvae, Chiapaphysa 202, 204 

guaraniana, Chilina 328 

guildfordiae, Tellina 232 

guineensis, Neilonella 68 

gussonii, Williamia 56 


haasi, Bostryx (Bostryx) 271 
Bostryx (Bostryx) minor 272 
Columbinia (Pfeifferiella) 271, 274 
Llaucanianus 271-272 
Naesiotus 271 
Systrophia (Systrophia) 271 
Hadoceras taylori 212 
Hadziella 204 
hainanensis, Brotia 300 
Haitia 204 
moreleti 207 
Halicardia flexuosa 81, 171 
halimera, Rhinoclama 80, 94-95, 122, 157, 
161 
Haliotidae 15, 77, 19, 19, 51 
Haliotis diversicolor aquatilis 15, 17, 19, 19, 
51 
Haloa japonica 16, 31, 31, 56 
Halonympha atlanta 80, 88, 97-98, 123, 
137,166 
atlantica 96 
depressa 80, 96, 123, 134, 153, 157 
Halopecten undatus 109 
Haminoeidae 16, 31, 37, 41, 45, 56 
Hamiota 304, 310 
subangulata 306, 308, 309, 311, 315, 
316-318 
hampsoni, Ledella pustulosa 69, 143, 159- 
160, 164 
Neilonella 68, 111, 113 
hanna, Yoldiella 70, 164, 167, 169-170 
Happia (Happia) florezi 270 
Harpa 254 
harpa, Physa (Costatella) 204 
harryi, Caribnauta 202, 204 
Hastula bacillus 55 
Hauffenia 204 
haustrum, Pomacea 353 
hayıi, Helix 227, 244 
Pedinogyra 244 
hedleyi, Dacrydium 72, 109-110 
Glacidorbis 55 
Heliacus variegatus 55 
Helicina 275 
(Trichohelicina) klappenbachi 272, 275 
Helicinidae 8, 53, 265, 272, 275 
Helicoradomenia juani 49 
Helix 235 


374 INDEX 


argellacea 245 
argilacea 245 
argillacea 245 
carmelita 245 
contermina 3, 4 
contusula 1-2, 5 
cunninghami 244 
fraseri 244-245 
hayıi 227, 244 
(Helicogena) contusula 4 
laevissimus 234 
mora 245 
Helminthoglypta alfi 176, 182, 201 
(Coyote) taylori 212 
Helminthoglyptidae 201, 212 
Hemicena cerrateae 269 
Hemisinus 235 
lineolata 235 
lineolatus 235 
hemphilli, Asthenotherus 78, 114 
Physella 204 
henriettae, Brotia 234 
Melania 234 
henrypilsbryi, Mesembrinus (Ornatimorus) 
271,274 
Mesembrinus (Ornatimormus) 
densestrigatus 270 
Mesembrinus (Ornatimormus) 
pichitacalugaensis 274 
herklotsi, Cyclophorus 15, 21, 22, 53 
hernandezae, Littoridina 271 
Hesperisternia vibex 239 
hessleri, Tindaria 68, 124, 153, 158-160, 173 
Hiatellidae 59, 228 
hickmanae, Choristella 51 
hilleri, Spinula 69, 94, 109, 112-113, 120, 
133, 142, 157,159, 164, 167-169, 173 
Hindsia 239 
acuminata 239 
nassoides 239 
Hipponicidae 15, 25, 27, 42, 54 
Hipponix 42 
comica 15.25: 27.54 
hispida, Coccopygya 52 
Histioteuthidae 349 
Histoteuthidae 349 
Horatia 204 
Horatiinae 204 
horrida, Laevicordia 81, 89-90, 97-98, 109- 
1197453 
hubbsi, Coahuilix 203, 205 
humboldtina, Physa (Costatella) 205 
humile, Linepithema 355 
hupensis, Onchomelania 54 
Hyalina 243 
Hyalopecten 73, 87, 98, 101-102, 113-114 


parvulinus 73, 126 
undatus 73, 99 
hydiana, Lampsilis 306, 308, 309, 310, 311, 
315, 316-318 
Hydrobiidae 54, 175, 185, 187-190, 194— 
196, 200-212, 265, 273-274 
hydroida, Thyasira 116 
hyltonscottae, Zilchogyra 271 
Hypselartemon 1-2, 3, 4, 8-10 
alveus 1-2, 3, 4, 6, 8-9 
contusulus 1-2, 3, 4-5, 9 
deshayesianus 1-2, 3, 4-6, 8-9 
paivanus 1-2, 3, 6, 6-/, 8-10 
Hypselodoris 44 
festiva 16, 32, 35, 56 
Hyriidae 229 


Idabasis 205 
idahoensis, Lymnaea 205 
Valvata 205 
Idasola 64 
iguazuensis, Chilina 321-322, 322-324, 
324-326, 326-328, 328-329 
Illex argentinus 347, 348, 349 
llyanassa obsoleta 341, 342 
imbricatus, Serpulorbis 15, 25, 27, 54 
imeldae, Bostryx (Elatibostryx) 270-271 
Bostryx (Elatibostryx) costifer 269 
imminens, Pyrgulopsis 205 
inaequisculpta, Bathyarca 71, 84, 99, 101, 
108-111, 113, 136, 140-142, 144, 149- 
152, 154, 158-161, 165-169 
incala, Yoldiella obesa 70, 125-129, 135, 
137, 139, 154-155 
Incania florezi 270 
Incertae cedis 81-82 
Incerte cedis 87-90, 94-99, 102-104, 107- 
108, 110-111, 114-117, 119, 122-125, 
128, 130, 136-138, 141, 143, 145, 147, 
150, 153, 160-161, 164-165, 169-173 
inconspicua, Yoldiella africana 70, 84, 159- 
16916265172 
Yoldiella inconspicua 70, 84, 87, 90-92, 
96-100, 102, 124, 127-128, 132, 135- 
138, 141-152 
Yoldiella profundorum 115, 117-120 
incrassata, Thyasira 86, 131, 134 
incrassatus, Axinulus 129-134, 137-139, 143 
Leptaxinus 74 
inflata, Cuspidaria 88, 98 
Thyasira 65, 75, 86, 96, 108-109, 118- 
120, 153,165, 1580178 
inflatiportus, Bostryx (Bostryx) obliquiportus 
271 
insculpta, Yoldiella 70, 84, 125-126, 129- 
134, 139, 156-158 


INDEX 375 


insoleta, Policordia 81, 93-95, 99, 121, 153, 
172 
insularum, Pomacea 299, 353 
intermedia, Limopsis cristata 72, 106-107 
intermedius, Thyasira 170 
intermontana, Radix 205 
Inversidens 308 
japanensis 305-306, 309, 311, 316-318, 
317-318 
iostoma, Cantharus (Prodotia) 238 
Pollia 238 
Triton 238 
iridella, Friersonia 312 
Ischnochiton comptus 15, 40, 49 
Ischnochitonidae 15, 40, 49 
isidroensis, Steeriana (Steeriana) 
celendinensis 272 


Jagora 336 
jaliscoensis, Amecanauta 201, 205 
jamesi, Ledella 68, 83, 107-108, 115-116, 
119 
Janthina janthina 54 
janthina, Janthina 54 
Janthinidae 54 
japanensis, Inversidens 305-306, 309, 311, 
316-318, 317-318 
Japeuthria ferrea 16, 27, 28, 54 
japonica, Assiminea 15, 24, 26, 54 
Asterophila 54 
Coceulina15,20:27,52 
Cypraea gracilis 44 
Haloa 16, 31, 31,56 
Nipponoscaphander 16, 31, 33, 56 
Siphonaria 16, 33, 35, 56 
Waldemaria 53 
javanica, Cryptosoma 237 
javanicum, Sinum 237 
jeffreysi, Cuspidaria 79, 93, 133, 156 
Policordia 81, 97, 142 
Yoldiella 70, 84, 87, 94, 97-102, 104, 
108-110, 118, 121, 123, 126-129, 131- 
133, 135-143, 149-150, 152-156, 161, 
168, 173 
johnsoni, Malletia 71, 84, 89-98, 100, 124- 
137, 140-141, 158-160, 165-167, 172 
juani, Helicoradomenia 49 
Juga 202, 205, 208 
acutifilos 202 
chrysopylica 202, 205 
junturae, Fluminicola 205 
Radix 205 
Juturnia 203, 205 


kansasensis, Biomphalaria 209 
Karevia 204 


Kelliella 65, 77, 112, 114, 116-117, 143, 148, 
150 
abyssicola 77, 108-110, 153, 160-161 
adamsi 77, 107 
atlantica 77, 86, 90, 97, 106-112, 117- 
120, 122, 124-133, 135-157, 159-160, 
163, 165-166, 168-170, 172-173 
biscayensis 77, 86, 121-123 
concentrica 77, 90-91 
elongata 77, 90, 106, 111, 115-116, 118- 
119, 146, 159, 170-171 
miliaris 77, 121, 128 
nitida 77, 96, 98, 100-101, 103 
tenina 77, 171 
Kelliellidae 59, 65 
klappenbachi, Helicina (Trichohelicina) 272, 
275 
knudseni, Amphiplica 51 
Cardiomya 79, 89, 96-98, 100, 115-117, 
128 
kolobensis, Fontelicella 205 
Pyrgulopsis 209 
kosteri, Tryonia (Paupertryonia) 205 
Kurilabyssia venezuelensis 51 


labio, Monodonta confusa 19, 20, 52 
Monodonta confusa forma 15 
lachayensis, Scutalus (Scutalus) versicolor 272 
lachrymosa, Scrippsiella 341-342, 342, 344— 
345 
laeta, Cipangopaludina chinensis 15, 17, 22, 
22100 
laeve, Cerithium 236 
Laevicordia 81, 95, 171 
horrida 81, 89-90, 97-98, 109-110, 153 
laevigata, Melania 234 
Laevipilina antarctica 49 
laevis, Cerithium 236 
Melania 234 
Poli¢ordia 81,112, 153 
laevissimum, Buccinum 256 
laevissimus, Helix 234 
Lambis plicata 238 
Lametila abyssorum 68, 83, 96-97, 99-100, 
113, 140, 142, 144 
Lametilidae 59 
laminifera, Limatula 72, 85, 92, 98 
Lamprotula leaii 229 
Lampsiline 303-304 
Lampsilis 304, 308 
hydiana 306, 308, 309, 310, 311, 315, 
316-318 
ovata 306, 308, 309, 310, 311, 315, 316- 
318 
straminea 306, 308, 309, 310, 311, 315, 
316-318 


376 INDEX 


lanceolata, Limopsis cristata 72, 164, 171 
lapidaria, Pomatiopsis 54 
laraosensis, Bostryx (Bostryx) obliquiportus 
202 
Temesa (Temesa) pilsbryi 272 
lasseni, Vorticifex 205 
lata, Yoldiella 70, 84, 124-137 
latecolumellaris, Naesiotus (Naesiotellus) 
272-273 
lateralis, Addisonia 51 
latestriata, Temesa (Temesa) 272 
lateumbilicatus, Radiodiscus 272 
Latiaxis mawae 241 
Laurentiphysa 205, 211 
chippevarum 202 
lavernensis, Gastrocopta (Gastrocopta) 205 
leaii, Lamprotula 229 
Unio 229 
Ledella 58, 69, 103-104, 155, 164 
aberrata 130, 151, 153=155, 173 
aberrenta 68, 100, 120 
acinula 68, 83, 106-107, 112, 152, 154, 
161,170 
acuminata 68, 83, 122-123, 137, 141 
galathea 154 
galatheae 68, 154, 163 
jamesi 68, 83, 107-108, 115-116, 119 
lusitanensis 68, 158, 160 
oxira 68, 112-113 
parva 69, 92 
pustulosa argentinae 69, 118 
pustulosa argentinea 83, 117-120 
pustulosa hampsoni 69, 143, 159-160, 
164 
pustulosa marshalli 69, 83, 136-140, 142- 
143, 145-150, 152 
pustulosa pustulosa 69, 83, 121-133, 
135-137, 139, 141, 143-144 
sandersi 69, 170-172 
similis 69, 123 
solidula 107 
sublevis 69, 83, 93-101, 115-117, 119, 
129-130, 134, 139-140, 142-144, 152, 
160, 162, 166 
ultima 63, 65, 69, 83, 87, 94, 98-105, 
109-113, 117, 119-120, 135, 142-145, 
147-155, 160-163, 165-169, 173 
Leguminaria costatus 231 
Lemiox 310 
rimosus 306, 309, 311, 315, 316-318 
lens, Myrtea 74, 105 
lenticula, Portlandia 71, 124, 132, 156, 159, 
162, 164 
Lepetellidae 51 
Lepetidae 50 
Lepetodrilidae 51-52 


Lepetodrilus 43 
nux 51 
pustulosus 51 
Lepidopleurus asellus 49 
Leptarionta woytkowskii 275 
Leptaxinus incrassatus 74 
Leptochitonidae 49 
Leptonidae 76, 114, 157, 172 
Leptomormus 272 
lesueurii, Ancistrocheirus 347 
Leucozonia tuberculata 240 
leve, Cerithium 236 
levior, Ampullaria 299 
Pomacea 299 
ligamentina, Actinonaias 306, 308, 309, 311, 
315, 316-318 
Lilloiconcha 272 
lima, Limalepeta 50 
Limalepeta lima 50 
Limatula 73, 88, 94 
bisecta 72, 121 
celtica 72, 105, 114, 142, 147,150, 152 
laminifera 72, 85, 92, 98 
louiseae 72, 88, 98, 102, 107, 111, 113- 
114, 123, 141, 147, 169, 173 
margaretae 73, 101, 107-108, 130, 137, 
142-146, 148-150, 152 
smithi 73, 164, 168-169, 171 
subovata 73, 85, 91-95, 117, 122-123, 
126-128, 130-138, 140-141, 149, 158 
limatula, Myonera 124 
Limatulidae 65 
Limea 73, 116 
argentineae 73, 115 
lirata 73, 85, 96 
sarsi 73, 85, 121 
Limidae 59 
Limifossoridae 49 
Limnaea (Stagnicola) albiconica 200 
(Pseudosuccinea) dineana 203 
Limopsidae 59, 65 
Limoposis 72,112, 115 
aurita 72, 85, 90, 121, 124 
cristata 85, 89-90, 94, 111, 116, 123, 138, 
164, 170 
cristata affinis 72, 91-93, 95 
cristata cristata 72, 123, 125, 156 
cristata lanceolata 72, 164, 171 
cristata intermedia 72, 106-107 
galathea 72, 85, 87, 99-103, 105, 109- 
110, 114, 152, 161, 163, 166, 168-169, 
1723 
galatheae 163, 169 
minuta 72, 157 
spicata 72, 85, 115-116 
surinamensis 72, 85, 105, 108-109 


INDEX 377 


tenella 65, 72, 85, 97-101, 113-114, 141, 
161-163, 167-168, 172 
lineata, Melania 235 
Pomacea 293-294, 299 
lineolata, Aylacostoma (Hemisinus) 235 
Melania 235-235 
lineolatus, Hemisinus 235 
Melanopsis 235 
Linepithema humile 355 
Lingulodinium polyedrum 345 
lirata, Limea 73,85, 96 
lischkei, Chlorostroma 52 
Lithoglyphinae 206 
Lithoglyphus 205-206 
Littoraria 228, 236 
(Littoraria) zebra 236 
pulchra 228, 236 
Littoridina 206 
hernandezae 271 
peiranoi 273 
similis 274 
Littoridininae 206-207, 209 
Littorina 45, 236 
brevicula 15, 17:23:25. 53 
pulchra 228, 236 
zebra 236 
Littorinidae 15, 172.23,25:53, 228; 236 
lividus, Toxolasma 306, 309, 310, 311, 315, 
316-318 
lizarasoae, Bostryx (Pseudoperonaeus) 272 
Llaucanianus haasi 271-272 
longicallis, Abra 77, 89, 162 
longispira, Bostryx (Pseudoperonaeus) 272 
Lopesianus 272 
crenulatus 270, 272 
lordi, Physa 201 
louiseae, Limatula 72, 88, 98, 102, 107, 111, 
113-114, 123, 141, 147, 169, 173 
Propeleda 69, 118-120 
lucida, Yoldiella 70, 84, 89-91, 121, 141 
lucidum, Parvamussium 73, 85, 106, 108, 
112-113 
Lucinidae 59, 74, 114, 166 
Lucinoma filosa 74, 88, 164, 169, 171-172 
lugoi, Mexipyrgus 206 
luhuanus, Strombus 15, 24, 26, 54 
lusitanensis, Ledella 68, 158, 160 
lusoria, Meretrix 290 
lutea, Tellina 232 
Tellina (Megangulus) 232 
Lutrilimnea 203, 206 
gentilis 204 
polyskelidis 206, 210 
ursina 211 
Luzonia simplex 80, 86, 125-126, 129, 135, 
158, 160, 164, 166, 171-172 


Lycoteuthidae 349 
Lycoteuthis 349 
Lyhnidia 204 
Lymnaea 183 
(Hinkleyia) pilsbryi 192 
idahoensis 205 
Stagnalis 16, 34, 35, 56 
(Stagnicola) mohaveana 207 
Lymnaeidae 16, 34, 35, 56, 183-184, 200, 
203-207, 210-211, 268 
Lyonsia 78, 90 
Lyonsiella 80, 115, 165, 171 
abyssicola 80, 89-93, 95, 100, 118, 122- 
129, 197 
formosa 80, 94, 113-114, 122, 156-158 
fragilis 80, 91-94, 100 
freilei 80, 158, 167 
perplexa 80, 89, 95, 97, 124, 153 
smidti 80, 94, 101, 105, 110, 153 
зибдиаагай 80, 112-113, 126, 153, 156 
Lyonsiidae 59 
Lyra 254 


macedoi, Scutalus (Vermiculatus) 272 
Mactra deaurata 231 
denticulata 231 
subtriangulata 231 
Mactridae 59, 78, 154 
maculata, Pomacea 293, 299 
Vis 255 
madagascariensis, Cyclostoma 237 
Tropidophora 237 
magellanicus, Placopecten 73, 88 
magus, Gibbula 232 
malita, Malletia 71, 105-107 
Malletia 71, 88, 133, 139 
abyssorum 71, 84, 87-88, 97-105, 110- 
111, 119-120, 139, 145-148, 150-155, 
158, 161, 169, 173 
cuneata 71, 84, 87, 96-102, 116, 118- 
119, 127-128, 134-136, 138, 140-151, 
153-157, 162 
grasslei 71, 105-106, 120 
johnsoni 71, 84, 89-98, 100, 124-137, 
140-141, 158-160, 165-167, 172 
malita 71, 105-107 
obtusa 71, 138, 141 
pallida 71, 84, 101-105, 112, 160-162, 
166-168, 173 
polita 71, 84, 87-88, 98, 102, 104, 109- 
112, 139, 149, 151-152, 154-155 
succisa atlantica 105 
surinamensis 71, 105-106 
Malletiidae 59 
mantaroensis, Temesa (Temesa) 
decimvolvis 272 


378 


maoria, Opimilda 55 
maranhonensis, Steeriana (Cylindronenia) 
terrestris 275 
Maranhoniellus 272 
marasensis, Scutalus (Vermiculatus) 
cuzcoensis 272 
margaretae, Limatula 73, 101, 107-108, 
130, 137, 142-146, 148-150, 152 
Marginellidae 55 
marianae, Pseudorimula 52 
Marisa cornuarietis 53 
marmorata, Physa 204 
marmoratus, Musculus 122 
marshalli, Ledella pustulosa 69, 83, 136- 
140, 142-143, 145-150, 152 
Martinela 8 
Mathildidae 55 
maura, Mitra 242 
Maurea 233 
mawae, Latiaxis 241 
Pyrula 241 
maximus, Thaumastus (Quechua) salteri 
272 
maximus, Pecten 290 
Mayabina 201, 206 
petenensis 209 
polita 209 
sanctijohannis 210 
tempisquensis 211 
mcalesteri, Silicula 71, 105, 114, 120 
megachlamys, Physa 206 
megalochlamys, Physa 200, 206 
megalocyathus, Enteroctopus 347 
Megannularia pulchra 237 
megastoma, Chilina 321, 323, 327-329, 329 
Melania 235, 335 
(Acrostoma) pisum 333 
(Balanocochlis) pisum 333 
bulbosa 208 
carolinae 233 
conica 234 
costula 233 
freethii 234 
frethii 234 
glans 333 
globulosa 234 
henriettae 234 
laevis 234 
laevigata 234 
lineata 235 
lineolata 235-236 
newberryi 208 
pisum 331-334, 333, 336 
quadriseriata 235 
retusa 235 
subcarinata 234 


INDEX 


Melaniidae 335 
Melanodrymia aurantiaca 52 
Melanoides 234 
tuberculata 301 
Melanopsis lineolatus 235 
Meleager 254-255 
Meleagris 255 
melina, Fontelicella (Natricola) 206 
Melo 227 
amphora 242 
broderipii 222 
georginae 222 
(Melocorona) broderipii 242 
miltonis 222, 242 
Melocorona 242 
Melongenidae 241 
Menetus carinifex 202 
micromphalus 207 
Meretrix lusoria 290 
meridionale, Propeamussium 74, 156 
Mesembrinus 272-273 
(Mormus) expansus altorum 268 
(Mormus) expansus orcesi 273 
(Ornatimormus) combinai 269 
(Ornatimorus) henrypilsbryi 271, 274 
(Ornatimormus) henrypilsbryi 
pichitacalugaensis 274 
(Ornatimormus) henrypilsbryi 
densestrigatus 270 
Mesodesma deauratum 231 
denticulata 231 
ornata 230 
solenoides 232 
subtriangulata 231 
Mesodesmatidae 231 
Mesoginella pygmaea 55 
metcalfi, Fontelicella 206 
Mexinauta 201, 206 
Mexipyrgus 190, 202, 206 
carranzae 202-203, 206-207 
churinceanus 202-203 
escobedae 203 
lugoi 206 
mojarralis 207 
multilineatus 207 
Mexithauma 206 
quadripaludium 206, 210 
Mexithaumatinae 206 
Microamnicola 207 
Microbeliscus 272-273 
micrococcus, Fontelicella (Microamnicola) 207 
Microdiscula ccharopa 55 
Microgloma 67, 82, 106-108, 112, 135 
pusilla 67, 87 
turnerae 67, 82, 108, 122-125, 127-128, 
135-137, 140-141, 152, 154, 157, 164 


yongei 67, 107-108, 135, 146, 151, 159, 
165-166 
micromphalus, Menetus 207 
Micropilina arntzi 49 
Micropilinidae 49 
microstriata, Aplexa 211 
Midotrochus beyrichii 52 
miliaris, Kelliella 77, 121, 128 
milleri, Cochliopina 207 
miltonis, Melo 222, 242 
Voluta 242 
minckleyi, Nymphophilus 207-208 
miniscula, Tindaria 68, 83, 98, 108, 118, 
154, 162, 167-169 
minor, Bostryx (Bostryx) haasi 272 
Similipecten 73, 170 
Steeriana (Steeriana) celendinensis 272 
Temesa (Temesa) decimvolvis 272 
minuta, Limopsis 72, 157 
Portlandia 71, 83, 163, 170 
Thyasira subovata 75, 163, 170 
minutissima, Cocculinella 51 
minutus, Promenetus 207 
mirabilis, Epiphragmophora 273 
Osteopelta 51 
mirandoi, Potamopyrgus 273 
mirolli, Physa 207 
Mitra chinensis 242 
maura 242 
orientalis 242 
Mitrella burchardi 54 
Mitridae 16, 77, 29, 30, 55, 242 
modestus, Bostryx (Bostryx) 
angelmaldonadoi 268 
Modiolarca tumida 76 
Modiolus 127 
mohaveana, Lymnaea (Stagnicola) 207 
mojarralis, Mexipyrgus 207 
Monodonta labio confusa 19, 20, 52 
labio forma confusa 15 
Montacuta 76, 88, 101, 149 
ovata 76, 122, 163 
Montacutidae 59, 141 
montrouzieri, Pisania 240 
mora, Helix 245 
Pleurodonte 245 
moreleti, Haitia 207 
multicarinata, Valvata 205 
Multifasciatus 273 
multiguttatus, Drymaeus (Ornatimorus) 273 
multilineatus, Mexipyrgus 207 
Murex vibex 239 
Muricidae 16, 17, 29, 30, 54, 241 
Musculus discors 72, 127 
marmoratus 122 
mustelina, Volvarina 55 


INDEX 329 


Mycetopodidae 229 
myojinensis, Shinkailepas 53 
Myonera 80, 86, 96, 103, 111-114, 117, 119, 
145, 158. 162, 165 
angularis 80, 101, 138, 140, 142, 154 
atlantica 80, 86-88, 92, 101, 109, 111, 
116-117, 126, 140, 147, 149, 155, 158, 
173 
demistriata 80, 92, 96, 98-99, 119, 141, 
158 
limatula 124 
octoporosa 80, 100-105, 113 
paucistriata 80, 86, 96, 100, 106, 116, 118, 
126 
tillamookensis 80, 170-171 
myopsis, Thracia 78, 93 
Myrtea lens 74, 105 
Mysella 77, 89-90, 93, 96, 99, 102, 111, 124, 
197, 152. 169, 163,165, 1701 
ovata 170 
tumidula 77, 126, 154 
verrilli 77, 88, 93-95, 98, 108, 115, 121, 
123-124. 128, 197) 163, 170 
Mytilidae 58-59, 72, 94, 152 
Mytiloidea 255 
Mytiloides 255 
Mytilus edulis 290-291 


Nacellidae 15-16, 17-18, 42—43, 45, 50 
Naeera 228 
Naesiotellus 273 
Naesiotus 272-273 
andivagus 268 
elegantulus 270 
gracillimus 271 
haasi 271 
(Naesiotellus) columellaris 273 
(Naesiotellus) latecolumellaris 272-273 
(Naesiotus) bambamarcaénsis 268 
(Naesiotus) bicolor 269 
(Naesiotus) geophilus 271 
(Naesiotus) subcostatus chamayensis 269 
(Maranhoniellus) fernandezae 270 
pilsbryi 274 
(Raphiellus) cerrateae 269 
(Raphiellus) turritus 275 
(Reclasta) tarmensis 275 
silvaevagus 274 
ziichi 275 
nanna, Paludestrina 210 
Nannobeliscus 272-273 
Nassa northiae 239 
reticulata 256 
Nassaria acuminata 239 
bitubercularis 239 
nassoides 239 


380 INDEX 


Nassariidae 16, 29, 29, 54, 228, 240 
Nassarius festiva 16, 29, 29, 54 
nassoides, Hindsia 239 
Nassaria 239 
Triton 239 
Natica bifasciata 236 
Naticidae 15, 24, 25, 42, 45, 54, 236 
natricina, Physa 200 
Physa (Haitia) 207 
Natricola 207 
Neaera 228, 232 
chinensis 228, 232 
Neara 228 
Neilonella corpulenta 68, 113 
guineensis 68 
hampsoni 68, 111, 113 
salicensis 68, 83, 91-97, 105-107, 109, 
122—130, 133-136, 148, 150, 155-156, 
158, 160, 162-165, 171 
seguenza 164 
whoii 68, 83, 87, 96-100, 104, 108-111, 
117-120, 142-150, 152-155, 157, 160- 
161, 164, 166-168, 170 
Neilonellidae 59 
Nenia (Columbinia) zischkai 276 
weyrauchi 276 
Neohoratia 204 
Neolepton 118 
profundorum 77, 86, 115-117 
Neoleptonidae 59 
Neomphalidae 52 
Neomphalus 43 
fretterae 52 
Neopetraeus arboriferus obesus 273 
arboriferus paucistrigatus 273 
camachoi 269 
weyrauchi 276 
Neopilina galatheae 49 
Neopilinidae 49 
Nephronaias 312 
Neraea 228 
Nerita albicilla 15, 17, 20, 21, 52 
Neritidae 15, 17, 20, 21, 52 
Neritiliidae 53 
Neroea 228 
chinensis 232 
nevadense, Sphaerium (Amesoda) 208 
nevadensis, Valvata 208 
newberryi, Melania 208 
Newboldius angiportus 268 
nipponica, Cocculina 52 
Nipponoscaphander japonica 16, 31, 33, 56 
nitens, Physa 206 
Pristigloma 67, 82, 87-88, 93, 95-104, 
109, 111, 113, 119, 134-138, 140-142, 
144, 150, 155, 157, 164, 167-169, 172 


nitida, Kelliella 77, 96, 98, 100-101, 103 
Thracia 78, 90, 93, 98, 117 
Niveotectura pallida 50 
nodulosa, Bentharca 71, 157 
Northia 239 
northiae 239 
northiae, Nassa 239 
Northia 239 
notabilis, Rhinoclama 80, 86-88, 96, 113, 
115, 123, 142, 144, 146, 149, 153 
Novaculina 230 
gangetica 231 
novaculina, Solen 230 
Nucinella pretiosa 67, 169 
Nucinellidae 59 
Nucula 152 
callicredemma 67 
Nuculana acuta 69, 88-89 
commutata 69, 120 
vestita 69, 83, 123, 160, 163, 165 
Nuculanidae 58-59, 140 
Nuculidae 59, 67, 114 
Nuculoidea bushae 67, 82, 88-90, 106, 122- 
130, 132-133, 135, 156, 158-159, 161, 
163, 165-169 
pernambucensis 67, 82, 106, 112 
Nuculoma callicredemma 113 
elongata 67, 105 
granulosa 67, 82, 90-95, 108, 123, 136, 
141 
granulose 164, 170 
perforata 67, 83, 114-116, 158-160, 163 
similis 67, 83, 89-94 
nux, Lepetodrilus 51 
Nymphophilinae 204, 208 
Nymphophilus 208 
minckleyi 207-208 


Obeliscus 272-273 
(Microbeliscus) silvaevagus 272, 274 
obesa, Cuspidaria 79, 86, 91, 95, 108, 128, 
130-131, 133-134, 139-140, 143, 147 
Yoldiella incala 70, 84, 125-129, 135, 
137, 139, 154-155 
Yoldiella obesa 70, 93-98 
obesus, Drymaeus (Diaphanomormus) 
coelestini 270, 273 
Drymaeus sulfureus 273 
Neopetraeus arboriferus 273 
Obliquaria 304, 310 
reflexa 306, 309 3101511:315:316-318 
obliquiportus, Bostryx (Bostryx) 273 
Bostryx (Bostryx) angispira 268 
Bostryx (Bostryx) inflatiportus 271 
Bostryx (Bostryx) laraosensis 272 
obliquum, Parvamussium 73, 108 


INDEX 381 


Obovaria 310 
olivaria 306, 309, 311, 315, 316-318 
rotulata 312 
obsoleta, llyanassa 341, 342 
Thyasira 75, 86, 88-89, 91, 95, 116, 122- 
127, 129-131, 133-134, 137, 139-140, 
159, 458 160 
obsolete, Thyasira 156-157, 163, 165 
obtusa, Cerithidea 235 
Malletia 71, 138, 141 
obvoluta, Systrophia (Systrophia) pilsbryi 274 
occidentalis, Thaumastus (Thaumastiella) 
273 
Thaumastus (Thaumastiella) debilisculptus 
270 
ockelmanni, Dacrydium 72, 85, 91-95, 115- 
117, 123-135, 137, 139-140, 156, 158, 
160-161 
octanglatum, Dentalium 15, 40, 50 
octoporosa, Myonera 80, 100-105, 113 
oculifera, Aplysia 16, 32, 33, 56 
Ohridohoratia 204 
Ohrigocea 204 
Oleacinidae 268-269 
olivaria, Obovaria 306, 309, 311, 315, 316- 
318 
Olivella borealis 55 
Olivellidae 55 
Olividae 243 
omissa, Temesa (Temesa) 273 
omissus, Scutalus (Vermiculatus) 273 
Onchidiidae 16, 17, 35, 36, 56 
Onchomelania hupensis 54 
onyx, Crepidula 15, 17, 23, 23, 53 
Opimilda maoria 55 
Orbitestellidae 55 
Orbitulites 255 
Orbulites 255 
orcesi, Mesembrinus (Mormus) expansus 
273 
Thaumastus (Thaumastus) 273 
Orcesiellus 273 
oregonensis, Fusitriton 15, 25, 26, 54 
Oregoniateuthis 349 
Oreobasis 208 
Oreoconus 208 
planispira 208-209 
Orientaliidae 208 
orientalis, Mitra 242 
Pectinodonta 50 
ormeai, Epiphragmophora 273 
ornata, Crassatella 230 
Eucrassatella 230 
Mesodesma 230 
Ornatimorus 273 
Orthalicidae 265, 268-276 


ortizi, Bostryx (Bostryx) 273 
ortizpuentei, Scutalus (Scutalus) 273 
Osteopelta 43 
mirabilis 51 
Osteopeltidae 51 
Ostracita 255 
Ostracites 255 
Ostreidae 59 
Otopoma 233 
ovata, Lampsilis 306, 308, 309, 310, 311, 
315, 316-318 
Montacuta 76, 122, 163 
Mysella 170 
Yoldiella 70, 84, 108 
Ovulidae 44 
owenii, Cyllene 222, 228, 240-241 
oxira, Ledella 68, 112-113 


Pachychilidae 233, 331-333, 335-337 
Pachychilus 234 
pachyostracon, Craterarion 182, 203, 208 
pacifica, Chiapaphysa 208 
paivana, Alcidia 6 
Streptaxis 6 
paivanus, Artemon 6 
Hypselartemon 1-2, 3, 6, 6-7, 8-10 
Streptartemon 6 
Streptaxis 6 
Streptaxis (Streptartemon) 6 
palizae, Solaropsis (Psadariella) 273-274 
pallida, Malletia 71, 84, 101-105, 112, 160- 
162, 166-168, 173 
Niveotectura 50 
Voluta 243 
Paludestrina nanna 210 
Paludina chinensis 233 
pulchra 233 
subcostata 233 
Paludiscala 208-209 
caramba 188, 202, 208 
Paludiscalinae 209 
Paludomidae 234 
Paludomus conica 234 
globulosa 234 
paludosa, Pomacea 353 
Pampasinus 273 
Pandora pinna 78, 121 
Panopea 228 
Paphies (Mesodesma) subtriangulata 
subtriangulata 231 
(Paphies) subtriangulatum 231 
papyracea, Periploma 78, 88 
Paracrostoma 332, 334, 336-337 
Paraplexa 201 
parva, Chilina 329 
Cuspidaria 79, 86-96, 99, 106, 113, 118, 


382 INDEX 


122-123, 125-142, 144-150, 153, 156- 
158, 160.166 
Ledella 69, 92 
Parvamussium 73, 87, 92, 99, 101, 109, 
112-113, 123, 126, 140, 160-161, 168 
lucidum 73, 85, 106, 108, 112-113 
obliquum 73, 108 
permirum 73, 142, 152 
parvulinus, Hyalopecten 73, 126 
Passer domesticus 355 
patagonica, Zygochlamys 73, 114 
Patella 42 
vulgata 50 
Patellidae 43, 45, 50 
paucistriata, Myonera 80, 86, 96, 100, 106, 
11611852126 
Propeleda 69, 163 
paucistrigatus, Neopetraeus arboriferus 273 
paupercula, Strigatella 55 
Paupertryonia 209 
paxillus, Strictispira 55 
pecos, Assiminea 209 
pecosensis, Fontelicella 209 
Pecosorbis 209 
Pecten maximus 290 
Pectinidae 58-59, 73, 94, 114, 119, 121, 
127, 152, 157-159, 164, 279 
Pectinodonta orientalis 50 
pectunculoides, Bathyarca 71, 85, 88-92, 
109, 121, 156, 169 
Bathyarca pellucida 170 
Peculator dedleyi 55 
Pedinogyra hayii 244 
peiranoi, Littoridina 273 
pellucida, Bathyarca pectunculoides 170 
Cornirostra 56 
Peltospiridae 52 
Pentadina 255 
perforata, Nuculoma 67, 83, 114-116, 158- 
160, 163 
Periploma papyracea 78, 88 
Periplomatidae 59 
permirum, Parvamussium 73, 142, 152 
pernambucensis, Nuculoidea 67, 82, 106, 
112 
Peronia verruculatum 16, 17, 35, 36, 56 
perplexa, Lyonsiella 80, 89, 95, 97, 124, 153 
Yoldiella 70, 84, 107 
perrieri, Tindaria 68, 102 
Peruinia albicolor 268 
flachi superba 275 
peruvianus, Pupoides (Pupoides) albilabris 
273 
petenensis, Mayabina 209 
peterseni, Zilchogyra 273 
Petrophysa 209 


Pfeifferiella 274 
Pharidae 230 
Phaseolidae 59 
Phaseolus 68, 82, 91, 95, 116, 120, 172 
Phenacolepadidae 53 
Phenacotaxus 270 
(Ataxellus) florezi 270 
Philine argentata 16, 31, 32, 56 
Philinidae 16, 37, 32, 41, 45, 56 
Pholadomyidae 59 
Phos billenheusti 238 
Phreatomenetus 209 
Physa 195 
acuta 208 
cisternina 206 
(Costatella) harpa 204 
(Costatella) humboldtina 205 
elata 201 
fontinalis 207 
(Haitia) natricina 207 
lordi 201 
marmorata 204 
mirolli 207 
megachlamys 206 
megalochlamys 200, 206 
natricina 200 
nitens 206 
skinneri 210 
Spiculata 206 
vernalis 205, 211 
Physella 209 
acuta 301 
hemphilli 204 
Physidae 175, 185, 195-197, 200-202, 
204—205, 207-211 
Physinae 204, 209 
pica, Turbo 255 
pichitacalugaensis, Mesembrinus 
(Ornatimormus) henrypilsbryi 274 
Pia so 1635S 
conica 351, 352-353, 354-355 
polita 353 
pilosus, Scutalus (Vermiculatus) 274 
pilsbryi, Drymaeus 274 
Lymnaea (Hinkleyia) 192 
Naesiotus 274 
Systrophia (Systrophia) obvoluta 274 
Temesa 274 
Temesa (Temesa) laraosensis 272 
Temesa (Temesa) primigenia 274 
Temesa (Temesa) shutcoénsis 274 
pinetorum, Fontelicella 205, 209 
pinna, Pandora 78, 121 
Pintadina 255 
Pisania 240 
crenilabrum 240 


INDEX 


fasciculatum 240 
montrouzieri 240 
Pisidium (Cyclocalyx) sanguinichristi 210 
Pisulina adamsiana 53 
pisum, Balanocochlis 333 
Melania 331-334, 333, 336 
Melania (Acrostoma) 333 
Melania (Balanocochlis) 333 
Sulcospira 331, 333, 333-334, 335-337, 
oof 
Placopecten magellanicus 73, 88 
planispira, Oreoconus 208-209 
Systrophia (Systrophia) 274 
Planorbella (Seminolina) wilsoni 211 
Planorbidae 200, 202, 205, 207, 209-211 
platyssima, Thyasira 95, 118, 123 
Plectomerus 303, 308, 310-312 
dombeyanus 306, 309, 311, 315, 316-318 
Plekocheilus 268, 273 
(Orcesiellus) tenuissimus 273, 275 
Pletholophus 229 
Pleurobema 310 
sintoxia 306, 309, 311, 315, 316-318 
Pleurobeminae 304 
Pleuroceridae 200, 202, 205, 208 
Pleurodonte mora 245 
Pleurodontidae 273-274 
Pleurostoma 255 
grandis 244 
Pleurotoma 255 
carinata 243-244 
crispa 244 
decussata carinata 244 
grandis 244 
speciosa 244 
Pleurotomariidae 52 
plicata, Amblema 306, 309, 311, 315, 316- 
318 
Lambis 238 
Thyasira excavata 75, 164 
plicatus, Strombus (Dolomena) plicatus 238 
Pliopholygidae 209 
Pliopholyx 209 
Plocameros 256 
Plocamoceros 256 
Policordia 81, 93, 106, 148, 154, 166, 171 
atlantica 81, 95, 128-129, 131-132, 138, 
147, 158, 160, 164 
densicostata 81, 91-93, 99, 136-137, 
171-172 
gemma 81, 99-101, 113, 118-119, 125- 
126, 128, 130, 132-135, 139, 143, 145- 
147,172 
insoleta 81, 93-95, 99, 121, 153, 172 
jeffreysi 81, 97, 142 
laevis 81, 112, 153 


383 


Polinices (Polinices) bifasciata 236 
polita, Malletia 71, 84, 87-88, 98, 102, 104, 
109-112, 139, 149, 151-152, 154-155 
Mayabina 209 
PHASS3 
Pollia 238 
iostoma 238 
polyedrum, Lingulodinium 345 
Polygyra (Erymodon) rexroadensis 210 
Polygyridae 210 
Polymesoda 196 
polymorpha, Dreissena 351 
polyskelidis, Lutrilimnea 206, 210 
Pomacea 293, 299, 351-352, 355 
bridgesii 353, 354 
canaliculata 293-294, 295, 296-297, 298, 
299-301, 351, 352-353, 354-355 
diffusa 299, 351, 353, 354 
dolioides 353 
gigas 299 
haustrum 353 
insularum 299, 353 
levior 299 
lineata 293-294, 299 
maculata 293, 299 
paludosa 353 
(Pomacea) pulchra 233 
scalaris 353 
Pomatiasidae 237 
Pomatiopsis lapidaria 54 
popeii, Popenaias 303-306, 308, 309, 311, 
314:915: 36-318 
Unio 304 
Popenaiadinae 304 
Popenaias 303-304, 308, 310-312 
buckleyi 304 
popeii 303-306, 308, 309, 311, 312, 315, 
316-318 
Poromya 78, 87-88, 106, 108, 113, 143, 
146-147, 168 
granulata 78, 110, 138 
tornata 78, 94, 98-99, 113-114, 134, 149, 
153,156 
Poromyidae 59 
Portlandia 107, 109 
abyssorum 71, 161, 167, 169, 173 
lenticula 71, 124, 132, 156, 159, 162, 164 
minuta 71, 83, 163, 170 
Potadoma 234 
freethii 234 
Potamides 255 
Potamididae 197, 202, 235, 237 
Potamis 255 
Potamopyrgue 210 
Potamopyrgus 210 
cheatumi 209 


384 


mirandoi 273 
Prelametila 68, 119, 168 

clarkei 68, 83, 97, 108, 119-120 
pretiosa, Nucinella 67, 169 
primigenia, Temesa (Temesa) pilsbryi 274 
Pristigloma 67, 105-106 

alba 67, 87, 95, 97-101, 103, 111, 113, 


119, 137, 141, 146, 152-153, 157, 167- 


168, 178 
nitens 67, 82, 87-88, 93, 95-104, 109, 
111, 113, 119, 134-138, 140-142, 144, 
150, 155, 157, 164, 167-169, 172 
Pristiglomidae 59 
pristis, Buccinum 239 
Prodotia 238 
profundorum, Abra 77, 86-88, 94, 103-110, 
123, 136, 142-154, 156-157, 165-166, 
168 
Neolepton 77, 86, 115-117 
Yoldiella 70 
Yoldiella inconspicua 115, 117-120 
Promenetus 209 
minutus 207 
umbilicatellus 209 
Propeamussidae 59, 64 
Propeamussium 74, 87, 94, 105-109, 112, 
117-118 
centobi 74, 157 
meridionale 74, 156 
thalassinum 74, 110 
Propeleda carpenteri 69 
louiseae 69, 118-120 
paucistriata 69, 163 
Propoleda carpenteri 83, 114—116 
protea, Tryonia 202, 205 
proteus, Flexipecten 73 
Flexopecten 121 
Protocuspidaria 79, 119, 149, 156 
atlantica 78, 92 
simplex 171 
simplis 78, 137, 157-158 
verityi 79, 87-88, 92, 112, 117, 144, 157, 
160, 166 
Provanna variabilis 53 
Provannidae 53 
Psadariella 274 
Pseudamussium clavatum 74, 121 
pseudobesus, Drymaeus 273 
Drymaeus (Mesembrinus) 270 
Pseudococculinidae 51 
Pseudoglandina 274 
agitata 268, 274 
pseudolata, Yoldiella 70, 84, 123-124, 135, 
150159 
Pseudoliva ancilla 55 
Pseudolividae 55 


INDEX 


Pseudoperonaeus 274 
Pseudorimula marianae 52 
Pseudotindaria 68, 140 
championi 68, 117, 119-120, 152 
erebus 68, 83, 100, 109-110, 113, 118, 
154, 161-162, 166-167 
Pseudotryonia 200 
Ptychobela griffithii 244 
Ptychobranchus 304, 308, 310 
fasciolaris 306, 309, 311, 315, 316-318 
Ptychodon 275 
(Unilamellatus) unilamellatus 275 
pubescens, Thracia 78, 122, 125, 128 
pulchella, Cinnalepeta 53 
pulcher, Turbo 237 
pulchra, Ampularia 233 
Cyclostoma 237 
Littoraria 228, 236 
Littorina 228, 236 
Megannularia 237 
Paludina 233 
Pomacea (Pomacea) 233 
Tropidophora 237 
pulchrum, Cyclostoma 237 
Punctoidea 212 
Pupillidae 200, 202, 204, 206, 210, 265, 268, 
273 
Pupoides (Ischnopupides) chordatus 
argentinus 268 
(Pupoides) albilabris peruvianus 273 
pusilla, Microgloma 67, 87 
Pusio 228, 238 
elegans 238 
pustulosa, Ledella argentinae 69, 118 
Ledella argentinea 83, 117-120 
Ledella hampsoni 69, 143, 159-160, 164 
Ledella marshalli 69, 83, 136-140, 142— 
143, 145-150, 152 
Ledella pustulosa 69, 83, 121-133, 135- 
137, 139, 141, 143-144 
pustulosus, Cyclopecten 74, 89, 106, 110, 
118 


Lepetodrilus 51 
pygmaea, Mesoginella 55 
Temesa (Temesa) albocostata 274 
Thyasira 75, 86, 89-92, 109, 116, 121- 
122, 168 
pygmaeus, Bostryx (Bostryx) 274 
Bostryx (Bostryx) costatus 269 
Pyrene testudinaria tyleria 240 
Pyrgulopsis 181, 187, 201-211 
blakeana 201 
cahuillarum 202 
cochisi 201 
confluentis 211 
imminens 205 


kolobensis 209 
sancarlosensis 201 
taylori 212 

Pyrula clavella 241 
mawae 241 
Striata 241 


quadrata, Verticordia 80, 95-96, 101, 109- 
110, 113-114, 128, 135-136, 149, 160- 
161: $60, 178 

quadripaludium, Mexithauma 206, 210 

guadriseriata, Melania 235 

quadritaeniatus, Drymaeus (Orodrymaeus) 
farrisi 274 

quadrula, Quadrula 306, 309, 377, 315, 
316-318 

Quadrula 310 
quadrula 306, 309, 311, 315, 316-318 
refulgens 306, 309, 311, 315, 316-318 


radiatus, Solen 231 
Radiocentrum taylori 212 
Radiodiscus florezi 270 
giganteus 271 
golbachi 271 
lateumbilicatus 272 
thomei 275 
wygodzinskyi 275 
Radix intermontana 205 
junturae 205 
Ranellidae 15, 25, 26, 42, 45, 54 
Rectartemon 1-2, 8 
(Hypselartemon) 2 
(Hypselartemon) alveus 1-2 
rectius, Rhabdus 50 
reflexa, Obliquaria 306, 309, 310, 311, 315, 
316-318 
refulgens, Quadrula 306, 309, 311, 315, 
316-318 
rehderi, Bostryx (Elatibostryx) 274 
reidi, Cerithidea 235-236 
Cerithium 236 
reticulata, Nassa 256 
retusa, Melania 235 
rexroadensis, Polygyra (Erymodon) 210 
Rhabdidae 50 
Rhabdus rectius 50 
Rhinoclama 162, 168 
abrupta 80, 137, 171 
halimera 80, 94-95, 122, 156, 161 
notabilis 80, 86-88, 96, 113, 115, 123, 
142, 144, 146, 149, 153 
Rhynchopelta concentrica 52 
rimosus, Lemiox 306, 309, 311, 315, 316- 
318 
Rissooidea 194, 273 


INDEX 385 


robusta, Fontelicella (Natricola) 207 
Thyasira 103, 118-119, 122 
Yoldiella 70, 84, 114, 116 

robustus, Scutopus 49 

rodeoensis, Goniobasis 202 
Cerithium 202 

rodriguezae, Bostryx (Bostryx) 274 

rosea, Euglandina 351 

rostrata, Cuspidaria 92 

roswellensis, Fontelicella 210 

rotulata, Obovaria 312 

rotundata, Glebula 306, 309, 311, 315, 316- 
318 

rudis, Voluta 243 


Sairostoma 8 
salicensis, Neilonella 68, 83, 91-97, 105- 
107, 109, 122-130, 133-136, 148, 150, 
155-156, 158, 160, 162-165, 171 
salteri, Thaumastus (Quechua) maximus 
272 
sancarlosensis, Pyrgulopsis 201 
sanctijohannis, Mayabina 210 
sandersi, Dacrydium 72, 85, 92, 96-98, 
112-114, 138-139, 143 
Ledella 69, 170-172 
sanguinichristi, Pisidium (Cyclocalyx) 210 
sapotalensis, Actinonaias 312 
sarcochrous, Bulimulus (Protoglyptus) 275 
sarsi, Limea 73, 85, 1 
Saulea vitrea 53 
Savaginius 210 
зауй, Solen 231 
scaevoscala, Gastrocopta (Gastrocopta) 210 
scalaris, Pomacea 353 
scheltemae, Spinula 69, 109-110, 117, 119— 
120 
Schizoplax 49 
Scolodontidae 265, 270-271, 274, 276 
scotophilus, Bostryx (Bostryx) 274 
Scrippsiella lachrymosa 341-342, 342, 344— 
345 
scripta, Euplica 16, 28, 28, 54 
Scrobicularidae 58-59 
Scutalus 275 
grandiventris 271 
(Scutalus) chiletensis 269 
(Scutalus) chiletensis granulatus 271 
(Scutalus) coraeformis debilisculptus 270 
(Scutalus) ortizpuentei 273 
(Scutalus) versicolor lachayensis 272 
(Vermiculatus) costulatus 269 
(Vermiculatus) cuzcoensis 270 
(Vermiculatus) cuzcoensis marasensis 
272 
(Vermiculatus) macedoi 272 


386 


(Vermiculatus) omissus 273 
(Vermiculatus) pilosus 274 
(Vermiculatus) culmineus zilchi 275 
Scutellastra flexuosa 50 
Scutopus robustus 49 
Scutus sinensis 51 
secunda, Bathyacmaea 50 
seguenza, Neilonella 164 
selectum, Calliostoma (Maurea) 232 
selectus, Trochus 232 
Selenoteuthis 349 
semiaperta, Epiphragmophora diluta 274 
semiclausa, Epiphragmophora diluta 274 
semiplicata, Bullaea 241 
Bullia 227, 241 
Sepioteuthis 256 
Sepiotheuthes 256 
Serapta 71, 148 
serpenticola, Taylorconcha 211 
Serpulorbis imbricatus 15, 25, 27, 54 
Shinkailepas myojinensis 53 
shotwelli, Carinifex 210 
shutcoénsis, Temesa (Temesa) pilsbryi 274 
Sibirenauta 201 


sieboldiana, Acusta despecta 16, 17, 36, 36, 


56 
Silicula filatovae 71, 84, 87, 96, 100, 102- 
105, 111, 123, 143-144, 146, 148-152, 
161, 166, 168-170 
fragilis 71, 97, 99-100, 118, 126, 128, 
133, 138, 141, 165 
mcalesteri 71, 105, 114, 120 
Siliculidae 59 
Siliqua 231 
costata 231 
silvaevagus, Naesiotus 274 
Obeliscus (Microbeliscus) 272, 274 
similes, Yoldiella 165, 168-169 
Similipecten 73 
minor 73, 170 
similis 73, 121 
similirus, Yoldiella 70, 105, 115, 118 
similis, Сугепа 228-229 
Ledella 69, 123 
Littoridina 274 
Nuculoma 67, 83, 89-94 
Similipecten 73, 121 
Venus 228 
Yoldiella 70 
simplex, Cyclopecten 74, 92, 105 
Luzonia 80, 86, 125-126, 129, 135, 158, 
160, 164, 166, 171-172 
Protocuspidaria 171 
simplis, Edentaria 103, 153 
Protocuspidaria 78, 137, 157-158 
simpsoni, Adipicola 72, 135 


INDEX 


Simrothiellidae 49 

sinaloae, Ultraphysella 210-211 

sinensis, Scutus 51 

Sinonovacula constrictus 230 

sintoxia, Pleurobema 306, 309, 311, 315, 


310-918 


Sinum javanicum 237 

sinuosa, Yoldiella 71, 107 
Sinupharus africanus 231 
sinusdulcensis, Tropinauta 210-211 
Siphonaria 45 


japonica 16, 33, 35, 56 


Siphonariidae 16, 33, 35, 56 

skinneri, Physa 210 

smidti, Lyonsiella 80, 94, 101, 105, 110, 153 
smithi, Limatula 73, 164, 168-169, 171 
smithii, Unio 230 

Solaropsis 274 


(Psadariella) palizae 273-274 


Solemya 67, 105-106, 108-109, 114-117, 


1227125. 165,167 
acherax 90 
grandis 67, 91-92, 99 


Solemyidae 59 


Solen 231 


africanus 231 
constrictus 230 
costatus 231 
novaculina 230 
radiatus 231 
sayii 231 
tenuis 231 


solenoides, Darina 232 


Erycina 231-232 
Mesodesma 232 


solidula, Ledella 107 

sonomae, Archiphysa 210 

souzalopesi, Drymaeus (Drymaeus) 274 
spathidophallus, Stenophysa 210 
speciosa, Gemmula 244 


Pleurotoma 244 


Sphaeriidae 208, 210 
Sphaerium (Amesoda) nevadense 208 
Sphaerospira 245 


fraseri 245 


spicata, Limopsis 72, 85, 115-116 
spiculata, Physa 206 
Spinula 69, 103-104, 112, 168 


filatovae 69, 83, 124, 126, 158, 160, 162, 
164 

hilleri 69, 94, 109, 112-113, 120, 133, 
142, 157, 159, 164, 167-169, 173 

scheltemae 69, 109-110, 117, 119-120 

subexisa 69, 83, 108, 132, 136-138, 140- 
he 140; 162 


Spiraxidae 268 


squamosus, Chiton 256 
stagnalis, Lymnaea 16, 34, 35, 56 
Steeriana 268 
(Cylindronenia) maranhonensis terrestris 
219 
(Steeriana) cajamarcana 269 
(Steeriana) celendinensis 269 
(Steeriana) celendinensis isidroensis 272 
(Steeriana) celendinensis minor 272 
Stenophysa 211 
spathidophallus 210 
Stenostylus zilchi 276 
Sternus vulgaris 355 
stocktonensis, Tryonia 211 
Stomatia 256 
straminea, Lampsilis 306, 308, 309, 310, 
311, 315, 316-318 
Streptartemon 8 
deformis 9 
paivanus 6 
Streptaxidae 1, 5, 8-9 
Streptaxis 1, 8 
alveus 2 
(Artemon) alveus 2 
crossei 6 
deshayesianus 5-6 
(Eustreptaxis) alveus 2 
(Eustreptaxis) deshayesianus 3 
paivana 6 
paivanus 6 
(Streptartemon) paivanus 6 
striata, Fusus 241 
Pyrula 241 
striatum, Taphon 241 
striatus, Fusus 241 
Strictispira paxillus 55 
Strictispiridae 55 
Strigatella paupercula 55 
zebra 16, 17, 29, 30,585 
Stromatia 256 
Strombidae 15, 24, 26, 42, 54, 237 
Strombus 42, 235 
campbelli 237-238 
deformis 238 
(Dolomena) plicatus plicatus 238 
(Doxander) vittatus campbelli 238 
luhuanus 15, 24, 26, 54 
Strophocheilidae 8 
subangulata, Erycina 231 
Hamiota 306, 308, 309, 311, 315, 316-318 
subcarinata, Melania 234 
subcircularis, Thyasira 75, 118, 125-127, 
129, 131, 158, 160, 164 
Yoldiella 71, 84, 87-88, 102-105, 110- 
111, 118, 142, 145-149, 152-155, 158, 
169, 169, 173 


INDEX 387 


subcostata, Paludina 233 
subcostatus, Naesiotus (Naesiotus) 
chamayensis 269 
Viviparus 233 
subequatoria, Thyasira 75, 116, 119-120, 
165, 167 
subequilateria, Yoldiella 84, 138, 141 
subexisa, Spinula 69, 83, 108, 132, 136— 
138, 140-141, 143, 149, 162 
sublevis, Ledella 69, 83, 93-101, 115-117, 
119, 129-130, 134, 139-140, 142-144, 
152,160; 162; 166 
subovata, Limatula 73, 85, 91-95, 117, 122- 
123, 126-128, 130-138, 140-141, 149, 
158 
Thyasira 112 
Thyasira atlantica 116 
Thyasira minuta 75, 163, 170 
Thyasira subovata 75, 86, 90-91, 117— 
119, 127-128, 131-132, 135-143, 147, 
149, 158-159, 164, 170-172 
subquadrata, Lyonsiella 80, 112-113, 126, 
153; 156 
Verticordia 162 
subrectum, Episiphon 15, 40, 50 
subterranea, Columbinia (Pfeifferiella) 275 
subtrangularis, Brevinucula 67, 112 
subtriangulata, Mactra 231 
Mesodesma 231 
Paphies (Mesodesma) subtriangulata 231 
subtriangulatum, Paphies (Paphies) 231 
subtrigonum, Epilepton 76, 138 
Subulinidae 8, 265, 272-274 
Succineidae 189 
succisa, Malletia atlantica 105 
Thyasira altlantica 75, 86, 89, 92, 96, 
106-108. BI #7; 4192 123, 9027183, 
169065, 470; 772 
Thyasira succisa 75, 86, 122-123, 125- 
127, 130, 134, 139-140 
Sulcospira 331-333, 336-337 
pisum 331, 333, 333-334, 335-337, 337 
Sulculus diversicolor aquatilis 51 
sulfureus, Drymaeus obesus 273 
sulurnalis, Turritella 238 
superba, Endodonta 275 
Peruinia flachi 275 
superbus, Austrodiscus tucumanus 272 
Bostryx (Multifasciatus) 275 
surinamensis, Limopsis 72, 85, 105, 108- 
109 
Malletia 71, 105-106 
susannae, Anodon 229 
Anodontites (Anodontites) exoticus 229 
Sutilizonidae 52 
suturalis, Columbella 240 


388 INDEX 


suturnalis, Turritella 238 

symbolicum, Campanile 236 

symmetros, Axinodon 76, 112, 125-127, 131, 
138, 142, 146 

Symphynota discoidea 229 

Systrophia (Scolodonta) eliseoduartei 270 
(Scolodonta) florezi 270 
(Systrophia) altorum 268 
(Systrophia) haasi 271 
(Systrophia) obvoluta pilsbryi 274 
(Systrophia) planispira 274 
(Systrophia) zilchi 276 

Systrophiidae 268 


tampicoensis, Cyrtonaias 306, 309, 311, 315, 
316-318 
Taphon 241 
striatum 241 
tarmensis, Naesiotus (Reclasta) 275 
Taylorconcha 211 
serpenticola 211 
taylori, Hadoceras 212 
Helminthoglypta (Coyote) 212 
Pyrgulopsis 212 
Radiocentrum 212 
Tellina guildfordiae 232 
lutea 232 
(Megangulus) lutea 232 
Tellinid 86 
Tellinidae 77, 95, 106, 135, 163, 170, 232 
Temesa (Neniatracta) adusta cuencaensis 
270 
(Neniatracta) bequaerti 268 
pilsbryi 274 
(Temesa) albocostata 268 
(Temesa) albocostata pygmaea 274 
(Temesa) decimvolvis 270 
(Temesa) decimvolvis crassicostata 269 
(Temesa) decimvolvis mantaroensis 272 
(Temesa) decimvolvis minor 272 
(Temesa) latestriata 272 
(Temesa) omissa 273 
(Temesa) pilsbryi laraosensis 272 
(Temesa) pilsbryi primigenia 274 
(Temesa) pilsbryi shutcoénsis 274 
(Temesa) zilchi 276 
Temnocinclis euripes 52 
tempisquensis, Mayabina 211 
tenella, Limopsis 65, 72, 85, 97-101, 113- 
114, 141, 161-163, 167-168, 172 
tenerum, Cochlodesma 78, 122, 128, 131, 
160 
tenina, Kelliella 77, 171 
tenuis, Anodon 229 
Solen 231 
Unio 229 


tenuissimus, Plekocheilus (Orcesiellus) 273, 
278 
Terebra 255 
africana 243 
variegata 243 
Terebridae 55, 243 
Teredinidae 59 
teres, Cuspidaria 157 
terrestris, Steeriana (Cylindronenia) 
maranhonensis 275 
testudinaria, Brotia 337 
Pyrene tyleria 240 
Thais clavigera 16, 17, 29, 30, 54 
thalassinum, Propeamussium 74, 110 
Thaumastiella 275 
Thaumastus 275 
(Quechua) salteri maximus 272 
(Scholvienia) weyrauch 276 
(Thaumastiella) occidentalis 273 
(Thaumastiella) occidentalis debilisculptus 
270 
(Thaumastus) orcesi 273 
thermalis, Fontelicella 211 
Thiara 335 
Thiaridae 235, 331-332, 335-336 
thomei, Bulimulus (Rhinus) 275 
Radiodiscus 275 | 
Thracia 78 iz 122128 1416517072 
conradi 78, 89, 91 
durouchouxi 78, 106 
gracilis 78, 122 
myopsis 78, 93 
nitida 78, 90, 93, 98, 117 
pubescens 78, 122, 125, 128 
Thraciidae 59 
Thyasira 75-76, 86, 88-89, 91-99, 101, 
103-104, 106-111, 113-117, 119-126, 
128, 135-138, 140, 142-144, 148-150, 
152-154, 156, 158, 163-167, 170, 173 
alleni 74, 85, 115, 160, 163-164, 169-171 
atlantica 74, 104, 145, 152 
biscayensis 74, 109, 152, 155-156, 172 
brevis 74, 85, 87, 93-94, 97-101, 104, 
117—119, 124-127, 129-135, 137-139, 
141-142, 145, 147, 149-153, 155, 162, 
164-168, 172 
bushae 74, 163 
carrozae 74, 85, 94, 115-117, 150, 158, 
163-165, 170-171 
croulinensis 74, 85, 88-98, 105-106, 114— 
116, 119, 121-122, 124, 136-138, 142, 144, 
147, 152, 159-161, 163-166, 170-172 
equalis 74, 87, 89-94, 97, 106, 114-115, 
117-119, 122-127, 130-131, 135, 139, 
141, 144-146, 148, 154-155, 158-159, 
161-164, 166, 170, 172 


INDEX 389 


eumyaria 75, 85, 116, 122-124, 158, 160, 
182 169171 
excavata plicata 75, 164 
ferruginea 75, 85, 87-99, 101, 103, 105- 
108, 114-122, 124, 126-127, 131-132, 
135-138, 140, 143-148, 150-152, 159- 
161, 165-166, 172 
hydroida 116 
incrassata 86, 131, 134 
inflata 65, 75, 86, 96, 108-109, 118-120, 
153.183, 188; 178 
intermedius 170 
obsoleta 75, 86, 88-89, 91, 95, 116, 122- 
127, 129-131, 133-134, 137, 139-140, 
155, 158, 160 
obsolete 156-157, 163, 165 
platyssima 95, 118, 123 
рудтаеа 75, 86, 89-92, 109, 116, 121- 
122,188 
robusta 103, 118-119, 122 
subcircularis 75, 118, 125-127, 129, 131, 
158, 160, 164 
subequatoria 75, 116, 119-120, 165, 167 
subovata 112 
subovata atlantica 116 
subovata minuta 75, 163, 170 
subovata subovata 75, 86, 90-91, 117- 
119, 127-128, 131-132, 135-143, 147, 
149, 158-159, 164, 170-172 
succisa altlantica 75, 86, 89, 92, 96, 106- 
108115, - 1171019) 423391525158, 463, 
170,172 
succisa succisa 75, 86, 122-123, 125- 
127, 130, 134, 139-140 
tortuosa 75, 86, 91-95, 109-110, 124, 
158-160, 164-165, 170-172 
transversa 75, 86-89, 99, 101, 103-105, 
107-118; 115-117, 118=120) 122. 142, 
144, 148, 152, 155, 157-158, 163-166, 
168, 170-172 
trisinuta 75, 88, 116 
ultima 75, 86, 125-126, 158-160, 162 
verrilli 75, 86, 95-96, 107-108 
Thyasiridae 58—59, 64-65 
tillamookensis, Myonera 80, 170-171 
Timoriena 256 
Timorienna 256 
Tindaria 68, 170 
agatheda 83, 156 
callistiformis 68, 83, 98-105, 109-111, 117, 
119, 126, 130, 143-144, 153, 161, 164, 
168-169 
hessleri 68, 124, 153, 158-160, 173 
miniscula 68, 83, 98, 108, 118, 154, 162, 
167-169 
perrieri 68, 102 


Tindaridae 59 
Tindariopsis 68, 107 
aeolata 68, 107-108 
agatheda 68, 107-107, 112 
toreuma, Cellana 50 
tornata, Poromya 78, 94, 98-99, 113-114, 
134, 149, 153, 158 
tortuosa, Thyasira 75, 86, 91-95, 109-110, 
124, 158-160, 164-165, 170-172 
Toxolasma 308, 310 
lividus 306, 309, 310, 311, 315, 316-318 
translucidus, Drymaeus (Drymaeus) 275 
transversa, Thyasira 75, 86-89, 99, 101, 
103-105, 107-113, 115-117, 119-120, 
122; 142,.144,148; 152, 155, 157-158; 
163-166, 168, 170-172 
triangularis, Astarte 88 
Goodallia 77 
Verticordia 80, 95, 99-101, 138, 141-142, 
144, 159, 166, 169 
tricarinata, Eglisia 238 
triccarinata, Amathina 56 
Trichohelicina 275 
tridentata, Zilchistrophia 275 
Tridonta elliptica 77, 121-122 
trigonus, Anodontites (Anodontites) 
georginae 229 
trisinuta, Thyasira 75, 88, 116 
Triton 239 
elegans 228, 238 
iostoma 238 
nassoides 239 
(Pusio) elegans 228, 238 
(Pusio) vexillum 240 
turbinelloides 239 
vexillum 240 
trivialis, Fontelicella 211 
Trochidae 15,53; 19; 20; 232 
Trochogyra 275 
Trochus bicarinatus 232 
cunninghami 232-233 
grayanus 232 
selectus 232 
Trophon 256 
Trophona 256 
Tropidomya 120 
abbreviata 80, 96, 110, 115-116, 118, 130 
diagonalis 80, 164 
Tropidophora articulata 237 
madagascariensis 237 
pulchra 237 
Tropinauta 201, 211 
sinusdulcensis 210-211 
truncatum, Cerithium 235-236 
Tryonia 203, 205, 209 
circumstriata 211 


390 INDEX 


gilae 204 
(Paupertryonia) adamantina 200 
(Paupertryonia) alamosae 200 
(Paupertryonia) brunei 202 
(Paupertryonia) kosteri 205 
protea 202, 205 
stocktonensis 211 
tschudii, Bulimus 269 
tubercularis, Turbinella 240 
tuberculata, Leucozonia 240 
Melanoides 301 
Turbinella 240 
tucumanus, Austrodiscus superbus 272 
tumida, Modiolarca 76 
tumidula, Mysella 77, 126, 154 
Turbinella tubercularis 240 
tuberculata 240 
turbinelloides, Triton 239 
Turbo pica 255 
pulcher 237 
zebra 228, 236 
turnerae, Microgloma 67, 82, 108, 122-125, 
127-128, 135-137, 140-141, 152, 154, 
157, 164 
turneri, Amoria 243 
Voluta 243 
Turridae 243 
Turris crispa 244 
Turritella sulurnalis 238 
Turritella suturnalis 238 
Turritellidae 238 
turritus, Naesiotus (Raphiellus) 275 
tylerae, Columbella 240 
tyleria, Pyrene testudinaria 240 
Tylomelania 336 


ultima, Ledella 63, 65, 69, 83, 87, 94, 98— 
105, 109-113, 117, 119-120, 135, 142- 
145, 147-155, 160-163, 165-169, 173 

Thyasira 75, 86, 125-126, 158-160, 162, 
165 
Ultraphysella 209, 211 
sinaloae 210-211 

umbilicatellus, Promenetus 209 

uncinata, Cavolinia 16, 32, 33, 56 

undata, Cuspidaria 79, 103 

undatus, Halopecten 109 

Hyalopecten 73, 99 

undosum, Buccinum 238 

Unilamellatus 275 

unilamellatus, Ptychodon (Unilamellatus) 275 

Unio 304 

childreni 229-230 
chilensis 230 
chinensis 230 
douglasiae 229 


leaii 229 

popeii 304 

smithii 230 

fenuls 229 
Unionidae 197, 203, 229, 303 
Urocoptidae 265 
ursina, Lutrilimnea 211 
Utahphysa 209, 211 


Valvata idahoensis 205 
multicarinata 205 
nevadensis 208 
Valvatidae 205, 208 
variabilis, Provanna 53 
variegata, Terebra 243 
variegatus, Heliacus 55 
Velata 256 
Velates 256 
veletta, Yoldiella 71, 125, 156, 159 
Velorita 228 
Vema ewingi 49 
Vemidae 49 
Veneridae 59, 78, 114, 171-172 
venezuelensis, Kurilabyssia 51 
ventricosa, Cuspidaria 79, 163 
Venus 229, 255 
similis 228 
Venustaconcha 310 
ellipsiformis 306, 309, 311, 315, 316-318 
veranyi, Bulimus 268 
verityi, Protocuspidaria 79, 87-88, 92, 112, 
1177144, 157%, 160; 166 
Vermetidae 15, 25, 27, 54 
Vermiculatus 275 
vernalis, Physa 205, 211 | 
verrilli, Brevinucula 67, 82, 95, 97, 99-100, 
102, 107-109, 113; 137, 1409442. 452, 
156-163 
Mysella 77, 88, 93-95, 98, 108, 115, 121, 
123-1245.128 87 168,478 
Thyasira 75, 86, 95-96, 107-108 
verruculatum, Peronia 16, 17, 35, 36, 56 
versicolor, Scutalus (Scutalus) lachayensis 
272 
Verticordia 80-81, 90, 95, 104, 110, 116, 
119, 121, 128, 136, 143, 160 
quadrata 80, 95-96, 101, 109-110, 113- 
114, 128, 135-136, 149, 160-161, 167, 
173 
зибдиаага 162 
triangularis 80, 95, 99-101, 138, 141-142, 
144, 159, 166, 169 
Verticordiidae 58-59 
vesicalis, Bulimulus (Bulimulus) angustus 268 
Vesicomyidae 59 
vestita, Nuculana 69, 83, 123, 160, 163, 165 


vexillum, Triton 240 
Triton (Pusio) 240 
vibex, Cantharus (Gemophos) 239 
Hesperisternia 239 
Murex 239 
vilchezi, Bostryx (Bostryx) 275 
Villorita 228 
virginica, Crassostrea 290, 344 
Vis 254-255 
maculata 255 
vitrea, Saulea 53 
vitreum, Dacrydium 72, 88-89 


INDEX 391 


Xanthonychidae 275 
Xylophaga 78, 122 
Xylophagidae 59 


Yaquicoccus 211 
Yaquicoccus bernardinus 201, 211 
Yoidiella 58, 71, 115, 118, 123, 136, 149, 
166 
americana 69, 83, 87-88, 100-105, 110- 
117, 120; 159 
argentinea 115-116 
argentinensis 69 


vitreus, Delectopecten 73, 85, 121, 123, 136 artipica 69, 162, 164, 166 

vittatus, Strombus (Doxander) campbelli 238 biguttata 69, 106-108, 112, 117 
Viviparacea 209 bilanta 69, 83, 158-160, 162, 167, 170- 
Viviparidae 15, 22, 22, 53, 233 172 

Viviparoidea 209 biscayensis 70, 83, 132, 134, 136, 139- 
Viviparus 233 140, 142-155 


chinensis 233 

subcostatus 233 
Voluta amphora 242 

auris-vulpina 244 

broderipi 242 

broderipii 242 

broderippii 242 

ferussacii 243 

georginae 242 

miltonis 242 

pallida 243 

rudis 243 

turneri 243 
Volutidae 242 
Volutomitra alaskana 55 
Volutomitridae 55 
Volvarina mustelina 55 
Vorticifex lasseni 205 
vulgaris, Sternus 355 
vulgata, Patella 50 


Waldemaria japonica 53 
Wanga 234 
wareni, Dacrydium 72, 92, 123, 135 
weyrauch, Thaumastus (Scholvienia) 276 
weyrauchi, Austroselenites 276 
Bostryx 276 
Bostryx (Platybostryx) 273 
Nenia 276 
Neopetraeus 276 
whoii, Neilonella 68, 83, 87, 96-100, 104, 
108-111, 117-120, 142-150, 152-155, 
157, 160-161, 164, 166-168, 170 
Williamia gussonii 56 
willinki, Bostryx (Bostryx) 275 
wilsoni, Planorbella (Seminolina) 211 
woytkowskii, Leptarionta 275 
wygodzinskyi, Radiodiscus 275 


blanda 69, 83, 117-120 

capensis 70, 83, 164, 167, 170-171 

curta 70, 83, 91-95, 106-107, 112, 115- 
116, 123-132, 134-136, 158-159, 164, 
166-167, 171-172 

dissimilis 70, 87, 99, 138, 141 

ella 70, 84, 87, 96-100, 108, 113, 118, 
129, 132, 142-150, 152, 154-155, 161, 
163, 167-168 

enata 70, 84, 87, 91, 108 

extensa 70, 84, 117 

fabula 70, 96-103, 107-109, 119-120, 
129, 136, 141-143, 145, 148-149, 151, 
155, 162-163 

fibula 146, 152, 167-169 

frigida 70, 84, 89-91, 108, 121 

hanna 70, 164, 167, 169-170 

inconspicua africana 70, 84, 159-160, 
162, 168; 172 

inconspicua inconspicua 70, 84, 87, 90— 
92, 96-100, 102, 124, 127-128, 132, 
135-138, 141-152 

inconspicua profundorum 115, 117-120 

insculpta 70, 84, 125-126, 129-134, 139, 
156-158 

jeffreysi 70, 84, 87, 94, 97-102, 104, 108- 
110, 118, 121, 123, 126-129, 131-133, 
135-143, 149-150, 152-156, 161, 168, 
173 

lata 70, 84, 124-137, 139 

lucida 70, 84, 89-91, 121, 141 

obesa incala 70, 84, 125-129, 135, 137, 
139, 154-155 

obesa obesa 70, 93-98 

ovata 70, 84, 108 

perplexa 70, 84, 107 

profundorum 70 

pseudolata 70, 84, 123-124, 135, 156, 159 


392 


robusta 70, 84, 114, 116 
similes 165, 168-169 
similis 70 
similirus 70, 105, 115, 118 
sinuosa 71, 107 
subcircularis 71, 84, 87-88, 102-105, 
110-111, 118, 142, 145-149, 152-155, 
198,168. 169173 
subequilateria 84, 138, 141 
veletta 71, 125, 156, 159 
Yoldiidae 58-59 
yongei, Microgloma 67, 107-108, 135, 146, 
151, 159, 165-166 


zebra, Littoraria (Littoraria) 236 
Littorina 236 
Strigatella 16, 17,29, 30, 55 
Turbo 228, 236 

zilchi, Bostryx (Bostryx) 275 


INDEX 


Bostryx (Bostryx) compactus 269 
Bostryx (Bostryx) glomeratus27 1 
Epiphragmophora 275 
Naesiotus 275 
Scutalus (Vermiculatus) culmineus 275 
Stenostylus 276 
Systrophia (Systrophia) 276 
Temesa (Temesa) 276 
Zilchiella 276 
grandiportus 271 
Zilchistrophia 276 
tridentata 275 
Zilchogyra 275-276 
cleliae 269 
franzi 271 
hyltonscottae 271 
peterseni 273 
zischkai, Мета (Columbinia) 276 
Zygochlamys patagonica 73, 114 


Vol. 
Vol. 
Vol. 
Vol. 
Vol. 
Vol. 
Vol. 
Vol. 
Vol. 
Vol. 
Vol. 
Vol. 
Vol. 
Vol. 
Vol. 


Publication dates 


40, No. 
41, No. 
41, No. 
42, No. 
43, No. 
44, Мо. 
44, No. 
45, No. 


45, No 


46, No. 
46, No. 
47, No. 
48, No. 


49, No 
49, No 


1 
1 


2 
1 
1 
1 
2 
1 
om 
1 
2 
1 
1 
at 
2 


—2 


—2 
—2 


—2 
—2 


17 Dec. 
22 Sep. 
31 Dec. 


18 Oct. 


20 Aug. 

8 Feb. 
30 Aug. 
29 Aug. 
22 Mar. 
23 Aug. 
30 Dec. 


20 July 


16 Feb. 
10 Nov. 


27 July 


VOL. 50, NO. 1-2 MALACOLOGIA 2008 


CONTENTS 


JOHN A. ALLEN 
Bivavia Of MeWeen anne se ee ee eee a ek in 5 


ANDRE Е. BARBOSA, VALDEMAR К. DELHEY & EUGENE V. COAN 
Molluscan Names and Malacological Contributions of Wolfgang Karl 
Weyrauch (1907-1970) with a Brief Biography .................. 265 


ANDRE F. BARBOSA, NORMA C. SALGADO & ARNALDO C. DOS SANTOS 
COELHO 
Taxonomy, Comparative Morphology, and Geographical Distribution of 
the Neotropical Genus Hypselartemon Wenz, 1947 (Gastropoda: 
Pulmonata!: Sires eae a Be Gd eed Dm à Foe wae se 1 


ERIC G. CHAPMAN, MARK E. GORDON, JENNIFER M. WALKER, BRIAN K. 
LANG, DAVID C. CAMPBELL, G. THOMAS WATTERS, JASON P. CUROLE, 
HELEN PIONTKIVSKA & WALTER R. HOEH 
Evolutionary Relationships of Popenaias popeii and the Early Evolution of 
Lampsiline Bivalves (Unionidae): Phylogenetic Analyses of DNA and 
Amino Acid Sequences from F and M Mitochondrial Genomes. ....... 303 


EE-YUNG CHUNG 
Ultrastructural Studies of Oogenesis and Sexual Maturation in Female 
Chlamys (Azumapecten) farreri farreri (Jones 8 Preston, 1904) 


(Pteriomorphia: Pectinidae) on the Western Coast of Когеа.......... 279 
EUGENE V. COAN 
Publication Dates of MALACOLOGIA Volumes and Issues .......... 361 


AUGUSTO C. CRESPI-ABRIL 
A New Record of the Presence of Two Spermatophoric Complexes in a 
Male Short-Fin Squid (Шех argentinus, Castellanos, 1960) .......... 347 


DIEGO E. GUTIERREZ GREGORIC & ALEJANDRA RUMI 
Chilina iguazuensis (Gastropoda: Chilinidae), New Species from Iguazu 


NaticnahPak-Aicemiithes awe yrs AA CRUE BAI fo rr S24 
ALAN R. KABAT & RICHARD |. JOHNSON 

Dwight Willard Taylor (1932-2006): His Life and Malacological Research. . 175 
SHIHO KATSUNO & TAKENORI SASAKI 

Comparative Histology of Radula-Supporting Structures in Gastropoda . . 13 


FRANK KOHLER, NORA BRINKMANN & MATTHIAS GLAUBRECHT 
Convergence Caused Confusion: On the Systematics of the Freshwater 
Gastropod Sulcospira pisum (Brot, 1868) (Cerithioidea, Pachychilidae) .. 331 


KING-LUN KWONG, PAK-KI WONG, SAM S. S. LAU & JAIN-WEN QIU 
Determinants of the Distribution of Apple Snails in Hong Kong two 
Decades after their ntallimyasion.., ...u.. wur ee CON, 293 


AGNETA PERSSON, BARRY C. SMITH, MARK S. DIXON & GARY H. WIKFORS 
The Eastern Mudsnail, /Iyanassa obsoleta, Actively Forages for, Consumes, 
and Digests Cysts of the Dinoflagellate, Scrippsiella lachrymosa........ 341 


RICHARD E. PETIT & EUGENE V. COAN 
The Molluscan Taxa Made Available in the Griffith & Pidgeon 
(1833-1834) Edition of Cuvier, with Notes on the Editions of Cuvier and 
ОВ WODA'S In0e TOREROS ik à sous a + ee we 219 
CHUONG T. TRAN, KENNETH A. HAYES & ROBERT H. COWIE 


Lack of Mitochondrial DNA Diversity in Invasive Apple Snails (Ampul- 
О ол gh deck td ds à ue Meg, a> колы à Sot 


MALACOLOGIA 


INTERNATIONAL JOURNAL OF MALACOLOGY 
Web: http://malacologia.fmnh.org 


SUBSCRIPTION AND PAST ISSUE ORDER FORM 


Name: 
Address: 
Personal rates: Per volume 
Subscription 
Regular $62.00 
Student discounted rate $32.00 
Single & past volumes $62.00 


Institutional rates: 


Subscription $82.00 
Single & past volumes $82.00 


Agency and dealership rates: 


Subscription $85.00 
Single & past volumes | $85.00 


Publication of MALACOLOGIA is irregular and may not coincide with the calen- 
dar year. Occasionally, more than one volume is published in the same year. 
Therefore, subscribers are invoiced by volume, not by year. Any changes in price 
will occur by volume, not by year. 


Each volume contains 2 numbers. Sometimes a volume is complete in one issue 
and sometimes each number is published separately. 


Subscriptions begin with the current volume. Current volume postage is included 
for subscribers having American or Canadian mailing addresses. Subscribers 
requiring overseas shipment of the journal are charged an additional $16.00 per 
volume ($8.00 per issue for 2 issues totaling some 400 to 600 pages and weigh- 
ing up to 4 lbs. 


Single and past volumes are available with the exception of volumes 17(1), 18 
and 28 which are out of print. Postage and handling for shipment to an American 
or Canadian address is $10.00 per volume, $5.00 per issue. For shipment to an 
overseas address it is $16.00 per volume, $8.00 per issue. Postage for orders 
weighing more than 4 Ibs. will be determined by weight and destination. 


Prepayment is required. Postal money orders and checks (US $'s) must be drawn 
on American banks. VISA and MASTERCARD payments are accepted for an 
additional processing fee of $2.00 per volume for individuals and $3.00 per vol- 
ume for institutions, agencies and dealerships. Electronic transfer of funds to a 
Malacologia bank account is also possible for an additional $10.00 fee. 


Address: Malacologia fax: (856) 854-0341 


P.O. Box 385 e-mail: malacolog@jersey.net 
Haddonfield, NJ 08033-0309 
U.S.A. 


MALACOLOGIA ADDRESSES 


BUSINESS—SUBSCRIPTIONS 


1°T CLASS MAIL INCLUDING AIR, CERTIFIED, REGISTERED, ETC.: 


MALACOLOGIA 

PO. Box 385 

Haddonfield, NJ 08033-0309 
U.S.A. 


e-mail: malacolog@jersey.net tel/fax: (856) 854-0341 


MANUSCRIPTS 


1°T CLASS MAIL INCLUDING AIR, CERTIFIED, REGISTERED, ETC.: 


MALACOLOGIA 

PO. Box 1222 

West Falmouth, MA 02574-1222 
U.S.A. 


e-mail: georgedavis99@hotmail.com tel/fax: (508) 457-0810 


MOL, 90. NO: 12 


10, 


11: 


INSTRUCTIONS FOR AUTHORS 


MALACOLOGIA publishes original research 
on the Mollusca that is of high quality and 
of broad international interest. Papers com- 
bining synthesis with innovation are particu- 
larly desired. While publishing symposia 
from time to time, MALACOLOGIA encour- 
ages submission of single manuscripts on 
diverse topics. Smaller papers of local geo- 
graphical or systematic content, and of high 
quality and interest, may be accepted as 
‘Research Notes’. Nearly all branches of 
malacology are represented in the pages 
of MALACOLOGIA. 

Manuscripts submitted for publication are 
received with the tacit understanding that 
they have not been submitted or published 
elsewhere in whole or in part. 
Manuscripts must be in English, but may 
include an expanded abstract in a foreign 
language as well as the usual brief abstract 
in English. Both American and British spell- 
ings are allowed. 

Unless indicated otherwise below, contribu- 
tors should follow the recommendations in 
the Council of Biology Editors (CBE) Style 
Manual. 

Be brief. 

Manuscripts must be typed on one side of 
good quality white paper, double-spaced 
throughout (including the references, tables 
and figure captions), and with ample mar- 
gins. Tables and figure captions should be 
typed on separate pages and put at the end 
of the manuscript. Make the hierarchy of 
headings within the text simple and consis- 
tent. Avoid internal page references (which 
have to be added in page proof). 

Choose a running title (a shortened version 
of the main title) of fewer than 50 letters and 
spaces. 

Provide a concise and informative abstract 
summarizing not only contents but also re- 
sults. A separate summary is not wanted. 
Supply between five and eight key (topic) 
words to go at the end of the Abstract. 

Use the metric system throughout. Micron 
should be abbreviated um. 

Illustrations are printed either in one col- 
umn or the full width of a page of the jour- 
nal, so plan accordingly. The maximum 
size of a printed figure is 13.5 x 20.0 cm 
(preferably not as tall as this so that the 
caption does not have to be on the oppo- 
site page). The figure captions are not to 
be part of the figure. List the figure cap- 
tions on a separate page. 


MALACOLOGIA 


12. 


15% 


14. 


ar 


16: 


E, 


2008 


Drawings and lettering must be dark black 
on white paper. Lines, stippling, letters, 
and numbers should be thick enough to 
allow reduction by 30% to 50%. Letters 
and numbers should be at least 2 mm (line 
art) or 3 mm (gray-step, color) high after 
reduction, but avoid letter sizes > 6 mm. 
Several drawings or photographs may be 
grouped together to fit a page. Photo- 
graphs are to be high contrast. High con- 
trast is especially important for histological 
photographs. 

All illustrations are to be numbered sequen- 
tially as figures (not grouped as plates or 
as lettered sub series), and are to be ar- 
ranged as closely as possible to the order 
in which they are first cited in the text. Each 
figure must be cited in the text. 

Scale lines are required for all none diagram- 
matic figures, and should be convenient 
lengths (e.g., “200 um”, not “163 um”). Mag- 
nifications in captions are not acceptable. 
All illustrations should be mounted, num- 
bered, labeled or lettered, i.e. ready for the 
printer. Be professional. Sloppy illustrations, 
labels, borders will not be accepted. If as- 
sistance is required of MALACOLOGIAS 
editorial staff, the author will be charged for 
the services rendered. All computer-gen- 
erated graphics must be submitted elec- 
tronically (e-mail or CD-ROM). Submit 
such graphics as TIFF or JPEG files. Line 
art (black and white) graphics must have 
a final resolution of 1200 dpi, gray-step 
and color graphics, 300 dpi. In line draw- 
ings, the minimum line width of 0.2 mm is 
required (as measured by final size). 

A caption should summarize what is shown 
in an illustration, and should not duplicate 
information given in the text. The caption 
must not be part of the figure but be pro- 
vided in a separate file that has all the figure 
captions listed in order. Each lettered ab- 
breviation labeling an individual feature in a 
figure must either be explained in each cap- 
tion (listed alphabetically), or be grouped in 
one alphabetic sequence after the Methods 
section. Use the latter method if many ab- 
breviations are repeated on different figures. 
Tables are to be used sparingly and verti- 
cal lines not at all. Horizontal lines are to be 
used only in the header and foot of the table. 
Submit all tables in a separate file. 


. References cited in the text must appear in 


the Literature Cited section and vice versa. 
Refer to a recent issue of MALACOLOGIA 
for bibliographic style, noting especially that 
serials are cited unabbreviated. Supply in- 


19. 


20. 


2 


— 


22. 


23. 


24. 


28, 


formation on plates, etc., only if they are 
not included in the pagination. 

In systematic papers, synonymies should 
not give complete citations but should re- 
late by author, date and page to the Litera- 
ture Cited section. 

For systematic papers, all new type speci- 
mens must be deposited in museums where 
other scientists may study them. Likewise, 
MALACOLOGIA requires that voucher 
specimens upon which a paper is based be 
deposited in a museum where they may 
eventually be re-identified. 


. Submission of manuscripts. Contact the 


editor (georgedavis99@hotmail.com) to de- 
termine the method of submission if the 
manuscript is > 40 pages and graphics have 
file sizes exceeding 2.0 MB or to obtain 
instructions as to where a ms is to be 
mailed if mailing is necessary. Send the ms 
files to georgedavis99@hotmail.com. Each e- 
mail should not exceed 3.0 MB. 

Very long manuscripts may require submis- 
sion of the manuscript in triplicate by mail. 
The second and third copies can be repro- 
ductions. Also submit with the paper copies 
a computer disk in Microsoft Word (PC ver- 
sion) containing the manuscript including all 
tables and illustrations. Amanuscript that has 
been revised must be re-submitted on com- 
puter disk in Microsoft Word (PC version). 
Authors who want illustrations returned 
should request this at the time of ordering 
reprints. Otherwise, illustrations will be main- 
tained for six months only after publication. 
An author’s address should include an e- 
mail address. 

Electronic submission of manuscripts. 
A. Covering e-mail 

Prepare an e-mail message addressed to 
the editor (georgedavis99@hotmail.com) with 
cc to tom.wilke@allzool.bio.uni-giessen.de. 
The message should provide all information 
necessary for manuscript submission. 

B. Manuscript files 

Manuscript files can be accepted in Microsoft 
Word format only (PC version). Name the 
files after the first author (e.g. SMITH.DOC). 
Keep the file size of graphics below 2 MB 
(save graphics as TIFF file with LZW com- 
pression or as JPEG file). Once the MS is 
accepted, uncompressed files have to be 
provided on CD-ROM. Provide the tables as 
a separate file. Attach all files to the cover- 
ing e-mail message. For any question re- 
garding electronic submission of graphics, 
contact the graphics editor at: 
tom.wilke@allzool.bio.uni-giessen.de 


26. 


ar. 


28. 


29. 


30. 


31. 


kk 


REPRINTS AND PAGE COSTS 


Reprints must be ordered from the Business 
Office at the time proof is returned to the copy 
editor. Later orders cannot be considered. When 
100 or more reprints are ordered, an author will 
receive 25 additional copies free of charge. 
PDF files of the paper for personal use are 
available with a purchase of reprints; free 
of charge for subscribers to Malacologia. Sub- 
scription must include current and 2 paid-for 
immediate past and/or future volumes. 
There is a U.S. $4.00 per word charge for 
each author’s change in page proof. 
Page costs must be paid prior to publication. 
Page cost recovery is required as follows. 
Regular subscribers:* 

Non-student: 10 pages free; U.S. $30.00 

for each additional page (EAP) 

Student™: 15 pages free; U.S. $20.00 EAP 
Non-subscribers: 

Non-student: 8 pages free; U.S. $60.00 EAP 

Student™: 15 pages free; U.S. $30.00 EAP 
A reduction or elimination of page charges 
may be possible under certain circum- 
stances by arrangement with the editor. A 
reduction of page charges may be negoti- 
ated for papers exceeding 30 pages. 
Color figures must be paid-for by the author. 
Costs are U.S. $300 for the first color page in 
a paper and U.S. $150 for each subsequent 
color page. 


SUBSCRIPTION PURCHASE 


Effective 1 July 2007, subscription rates per 
volume are as follows: 

Personal: U.S. $62.00 

(Student discounted rate: U.S. $32.00) 

Institutional: U.S. $82.00 

Agency and Dealership: U.S. $85.00 
Current volume postage is included for ship- 
ment to an American or Canadian address. 
Overseas postage is $8.00 per issue; $16.00 
per volume. Single and past volumes are 
available at subscription rates. Postage and 
handling is $5.00 per issue; $10.00 per vol- 
ume for shipment to an American or Cana- 
dian address. Overseas postage is as cited 
above. Postage for orders weighing over 4 
lbs. will be determined by weight. 


Regular subscribers are those who have paid- 


up subscriptions for the current volume and two 
immediate past and/or future volumes. 
Students must identify themselves at the time 
of manuscript submission and also provide the 
e-mail address of their advisor. 


SMITHSONIAN INSTITU 


DO iit 


VOL. 50, NO. 1-2 MALACOLOGIA 


CONTENTS 
FULL PAPERS 


ANDRE Е. BARBOSA, NORMA С. SALGADO 8 ARNALDO С. DOS SANTOS 
COELHO 
Taxonomy, Comparative Morphology, and Geographical Distribution of 
the Neotropical Genus Hypselartemon Wenz, 1947 (Gastropoda: 


РУО ИЕ: SIMMS) he ee es 1 
SHIHO KATSUNO & TAKENORI SASAKI 

Comparative Histology of Radula-Supporting Structures in Gastropoda . . 13 
JOHN A. ALLEN 

EN OO A ........ nn au use du 57 
ALAN В. KABAT & RICHARD I. JOHNSON 

Dwight Willard Taylor (1932-2006): His Life and Malacological Research .. 175 


RICHARD E. PETIT & EUGENE V. COAN 
The Molluscan Taxa Made Available in the Griffith & Pidgeon 
(1833-1834) Edition of Cuvier, with Notes on the Editions of Cuvier and 
on Woods Index TOSTACECIOGICUS .......... nen ur 219 


ANDRE F. BARBOSA, VALDEMAR K. DELHEY & EUGENE V. COAN 
Molluscan Names and Malacological Contributions of Wolfgang Karl 
Weyrauch (1907-1970) with a Brief Biography .................. 265 
EE-YUNG CHUNG 
Ultrastructural Studies of Oogenesis and Sexual Maturation in Female 
Chlamys (Azumapecten) farreri farreri (Jones & Preston, 1904) 
(Pteriomorphia: Pectinidae) on the Western Coast of Korea. ......... 279 


KING-LUN KWONG, PAK-KI WONG, SAM $. $. LAU & JAIN-WEN QIU 
Determinants of the Distribution of Apple Snails in Hong Kong two | 
Decades after their Initialinvasion‘. . . . . ....,...,.,.,......... 293 
ERIC G. CHAPMAN, MARK E. GORDON, JENNIFER M. WALKER, BRIAN K. 
LANG, DAVID C. CAMPBELL, G. THOMAS WATTERS, JASON P. CUROLE, 
HELEN PIONTKIVSKA & WALTER R. HOEH 
Evolutionary Relationships of Popenaias popeii and the Early Evolution of 
Lampsiline Bivalves (Unionidae): Phylogenetic Analyses of DNA and 
Amino Acid Sequences from F and M Mitochondrial Genomes........ 303 


RESEARCH NOTES 


DIEGO E. GUTIERREZ GREGORIC & ALEJANDRA RUMI 
Chilina iguazuensis (Gastropoda: Chilinidae), New Species from Iguazu 
A A 321 


FRANK KOHLER, NORA BRINKMANN & MATTHIAS GLAUBRECHT 
Convergence Caused Confusion: On the Systematics of the Freshwater 
Gastropod Sulcospira pisum (Brot, 1868) (Cerithioidea, Pachychilidae) .. 331 


AGNETA PERSSON, BARRY C. SMITH, MARK S. DIXON & GARY H. WIKFORS 
The Eastern Mudsnail, IIyanassa obsoleta, Actively Forages for, Consumes, 
and Digests Cysts of the Dinoflagellate, Scrippsiella lachrymosa. . ...... 341 


AUGUSTO C. CRESPI-ABRIL 
A New Record of the Presence of Two Spermatophoric Complexes in a 
Male Short-Fin Squid (Шех argentinus, Castellanos, 1960) .......... 347 


CHUONG T. TRAN, KENNETH A. HAYES & ROBERT H. COWIE 
Lack of Mitochondrial DNA Diversity in Invasive Apple Snails (Ampul- 
I oe hte sa ey hn AAA 351 
LETTER TO THE EDITOR 
EUGENE V. COAN 
Publication Dates of MALACOLOGIA Volumes and Issues .......... 361 


INDEX 363 


x sa QE $ 


AT >» 


aa 


[RL L 


ra 
a> 
1: 


YAN 
Pte 
Sr 
de 


RAN 
«зах 
(ER 


23e 
ATEN 

u 
ae 


LE NO En 
y 
Pres : A 
5 RER NE \ 
L'EST > i \ 
RY ns ` ret 
ces iy vis 
PPL oe pe AN MERI OW CR aoe у т: 
: in Veste El N Pers 
| y ÓN 4 \ 
ones y UGG gece 
ares “ } \ 
SET Een i Е АН ВН i hat 
и LC ES Mase PAST) i TAOS Misses i Poda \ 
ER ality Gene 
ARR \ ‘ « er 
ei Sige 
eue О ИР 
м ИИ 
OPP ОО a uhr 
ton Tante, LE 
VERTUS N ; 
Lary abn ‘ ; en 


DORA 


ner мк 
ТУ Ел 


VER + 
А eget ns 


vice 


лить 


Pr AOS SP Meg 
ASR ER es 


о gO Se ad = 
Verne. BS 


Ань с 


nur 


PUES UE ROME, Sus 
LDLC ; 


Ware veg tes 
y IN 


CC 
IA 


IES TES 


Risks beans 


Яны 
аа bie aise 


pars tee 
сия ув 


u $ 
PR FT 


им» 


Зе, чм 


Peres, 


ETAT TRES 


ONE 
аи: 


Pere 
ОА 
ть а 


Peon gee ae, 
PENS haut, 
А, 


CEST 


NOUS 
6 кА 
HO TES : 

(EYES 


Sat бан 


Lee CERCA oe 
BSS OR PE tay ie 


CRE 
TANT 


y 


зе 
Фуке: 


O 
UA a 


EA На 


Var 


he Sew i eT RSG 
RTE TOI phy oye 
.. м 


ях: 


Paina: cae Wier 
PETER ENST ER * 


АА 
Hy ENTE ins 


& 
.. 


‘ 
‘+ 
\
Internet Archive Audio

Featured

Top

Images

Featured

Top

Software

Featured

Top

Texts

Featured

Top

Video

Featured

Top

Mobile Apps

Browser Extensions

Archive-It Subscription

Save Page Now

Full text of "Malacologia"

See other formats
